Hawaiian Plant Life: Vegetation and Flora 9780824846695

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Hawaiian Plant Life

PUBLICATION OF THIS BOOK IS SUPPORTED BY GRANTS FROM

Figure Foundation AND THE

Smithsonian Institution

Hawaiian Plant Life V EGETATION and FLORA  ROBERT J. GUSTAFSON DERRAL R. HERBST PHILIP W. RUNDEL

UNIVERSIT Y OF HAWAI‘I PRES S HONOLULU

PUBLICATION OF THIS BOOK WAS MADE POSSIBLE BY GENEROUS DONATIONS FROM THE FOLLOWING ORGANIZATIONS AND INDIVIDUALS

Figure Foundation Smithsonian InstitutionMuseum of Natural History Publication Fund and the Walcott Botanical Publication Fund University of California Los Angeles, M. Stanley Rundel Fund Laurel Woodley Published with the support of the Maurice J. Sullivan & Family Fund in the University of Hawai‘i Foundation.

© 2014 University of Hawai‘i Press All rights reserved Printed in China 19 18 17 16 15 14   6 5 4 3 2 1

Library of Congress Cataloging-in-Publication Data Gustafson, Robert J., author. Hawaiian plant life : vegetation and flora / Robert Gustafson, Derral R. Herbst, and Philip W. Rundel. pages cm Includes bibliographical references and index. ISBN 978-0-8248-3710-5 (cloth : alk. paper) 1. Plants—Hawaii. I. Herbst, Derral R., author. II. Rundel, Philip W. (Philip Wilson), author. III. Title. QK473.H4G87 2014 581.9969—dc23 2014004818

University of Hawai‘i Press books are printed on acid-free paper and meet the guidelines for permanence and durability of the Council on Library Resources. Designed by Mardee Melton Printed by Regent Publishing Services

CONTENTS

vii

Foreword

ix

Notes on the Photogra phy

xi

Preface

1

1. LANDFORMS AND CLIMATE

19

2. PLANT DIVERSITY AND EVOLUTION

33

3. PLANT COMMUNITIES

49

4. FLOWERING PLANTS

193

5. FERNS AND FERN ALLIES

225

6 . P O LY N E S I A N I N T R O D U C T I O N S

233

7 . A L I E N I N VA D E R S

243

8 . C O N S E R VA T I O N O F T H E H A W A I I A N F L O R A

255

Appendix

289

Glossar y

293

Fur ther Reading

309

Photogra ph Credits

311

Index

Anectochilus sandvicensis

FOREWORD

an inviting introduction designed to pique curiosity, not to quell it. You will want to learn more. Learning more about threatened and endangered plants is essential to conserving them. We want to try to save plant species of interest, such as the silverswords, not only because they are striking in appearance but because they represent a series of unique adaptations that have happened only in the Hawaiian Islands. People try to save only what they know about. Thus, a book that presents iconic images and stories of distinctive features of Hawaiian plants, as this book does, helps to lead the way in conservation efforts. Most people who have at least some knowledge of Island plants know the Hawaiian flora as the world’s most interesting oceanic island assemblage because of the isolation, relatively large land area, diverse climates from very wet to very dry and hot to cool, and because geological time has elapsed so that distinctive plant adaptations have been able to occur. The Hawaiian flora is also the most endangered island flora of major consequence in the world. Hawaiian plants have evolved in environments free from predators and invasive mainland species. With the introduction of the pests and invaders, the inevitable is happening. Hawaiian Plant Life: Vegetation and Flora is the first thoroughgoing attempt to record a large segment of native species in color photographs, and it may well be the last great photographic documentation of a fascinating flora. Some plants depicted are known from only one or a few individuals, and some are now extinct in the wild. Consequently, it is not just another book about Hawaiian plants; it is a unique one. It should be treasured not as an addition to a library on island organisms, but as a way of showing how plants evolved on the Hawaiian Islands, the products of evolution laid out like a banquet of biodiversity. What remains on the Islands is well worth studying, and many stories remain to be elucidated. But the enterprise of these authors in wanting to present visual records of a wonderful and waning flora is to be congratulated. It is a document at a unique moment in time, a document of Island biology that will not be replaced or superseded.

Visual contact with a flora is essential for becoming acquainted with the plants and what is significant about them. Thus, Hawaiian Plant Life: Vegetation and Flora becomes of prime importance in introducing Hawaiian native plants to a wide audience, both amateur and professional. Its wide coverage gives the newcomer to the Hawaiian flora an opportunity to see some of the remarkable adaptations that exist. The shapes of flowers, for example, in native Hibiscadelphus, Hibiscus, Pisonia, Metrosideros, and above all the lobeliads (Clermontia, Cyanea, Trematolobelia), show that the template evolved to suit bird pollination by native honeycreeper finches and the meliphagids. But the shapes without colors are only part of the story, and the value of this book is that it shows both. One can visualize the colors and positioning of these flowers as lures for those magnificent birds. At the other end of the spectrum, one can see the inconspicuous white tubular flowers, pollinated by insects and perhaps even beetles in some cases. Hawaiian plants have the floral colors and shapes of a flora poor in long-tongued bees, in contrast to a continental flora. Instead of bees, nectar-seeking flies and moths may have influenced the way that Hawaiian plants evolved. The least colorful flowers are, predictably, those of wind-pollinated plants, such as sedges and grasses—flowers that work well in an environment poor in pollinating insects. This subject of pollination biology and evolution of the flora is one that richly deserves study. The above examples are intended not as a scientific lesson but as an example of what books can do by way of inspiration. Sizes, shapes, and colors of flowers and other plant parts are not random; they are closely keyed to which plants were able to arrive by long-distance dispersal over a period of perhaps 10 million years and the ecological opportunities with which they were confronted. These are the stories of immigrants and how they changed over time. One recent large-format book with full-page color photographs of some rare Hawaiian plants told nothing about the significance of this amazing flora. By emphasizing “pretty” over evolutionary stories, that book conveyed the hidden message that there is nothing to learn—or nothing that can be told to an average person. The selection of pictures in that book emphasized only the more colorful flowers, as though it was a flower shop with items for sale. In contrast, Hawaiian Plant Life: Vegetation and Flora does not wall off the many stories to be told in the Hawaiian flora from viewers and readers. Instead it is

SHERWIN CARLQUIST

Santa Barbara, California

vii

Spermolepis hawaiiensis

NOTES ON THE PHOTOGRAPHY

My interest in the Hawaiian flora began in earnest during the mid-1970s when I first came to photograph Hawaiian native plants. The focus in the beginning was on rare and endangered species. But by 1980 I had decided to broaden my efforts to include as many endemic and indigenous plant species as possible. The photographs were taken over a span of eighteen years, with many trips back and forth to the Islands. Almost all of the plants pictured in this book were photographed in the field, with a few in botanical gardens or university collections. Nevertheless, some rare plants always remained elusive. It is with special gratitude that I acknowledge Tim Flynn, Angela Kay Kepler, and John Obata for their assistance in the field. I owe a debt of thanks to Gerry Carr for making a number of photographs available for this book from his collection. Much support to make this project possible came from Don Reynolds and the Natural History Museum of Los Angeles. Other individuals who contributed pictures are acknowledged elsewhere. I will always be grateful to the people listed at the end of the preface for their contributions and willingness to help with this project. Without their assistance and patience, this book would never have been possible. While many Hawaiian plants are on the verge of extinction today, rare new species are still being discovered thanks to intrepid botanists such as Steve Perlman, Ken Wood, Bob Hobdy, and others to mention only a few. Hawaiian plants are truly unique and occupy a special and significant place in the world’s floras. May they continue to survive.

ROBERT J. GUSTAFSON

ix

Lysimachia hillebrandii

PREFACE

The publication of the monumental Manual of the Flowering Plants of Hawai‘i (hereafter often referred to simply as the Manual) by the University of Hawai‘i Press in 1990 was a transformative accomplishment in providing the first complete and modern systematic treatment of the remarkable flora of the Hawaiian Islands. This Manual has done much to stimulate a renewed interest in the diversity and evolutionary origin of this flora among professional botanists. However, the Manual’s technical format and imposing length of 1,853 pages, split between two volumes, has limited its accessibility and use by amateur botanists and others interested in the practical identification of the native flora. Moreover, ferns and fern allies, which are common in the Hawaiian Islands and comprise nearly 15% of the Hawaiian flora, were not included in this Manual. This gap in the nonflowering vascular plants was filled with the publication of Hawai‘i’s Ferns and Fern Allies by Daniel D. Palmer (University of Hawai‘i Press, 2003). As with the Manual, however, this is a highly technical systematic treatment and not very user-friendly for readers who lack specialized training in plant systematics. In addition, illustrations in both the Manual and Fern Flora volumes are limited to a small number of line drawings. For many years we have heard from lay botanists, tropical gardeners, ecologists, and others interested in Hawaiian natural history that there is a strong need for a book to fill the gap between the small-format picture book represented by Plants and Flowers of Hawai‘i (Sohmer and Gustafson, University of Hawai‘i Press, 1987) and the technical floristic manuals of Hawaiian plants. This would be a book that would provide color photographs of a substantial portion of Hawaiian species, including all of the native genera, as well as include a concise description of each genus and the species illustrated. Thus, it could be used either as a stand-alone volume or as a companion volume to Manual of the Flowering Plants of Hawai‘i and Hawai‘i’s Ferns and Fern Allies. This has been our goal in developing this book project. The objective of Plant Life of Hawai‘i: Vegetation and Flora is to provide an illustrated guide to the Hawaiian native flora and vegetation, with high-quality color photographs of virtually all genera and more than half of the vascular plant species. Our book includes eight chapters. Chapter 1 describes landforms and climate in the Hawaiian Islands, including the Northwestern

Hawaiian Islands, and provides a general perspective on the physical geography of the archipelago. Chapter 2 offers an overview of the flora of flowering plants and ferns and the evolutionary processes that have led to this unique flora. Chapter 3 summarizes the vegetation of the Islands. Chapter 4 on flowering plants and chapter 5 on ferns and fern allies provide the illustrated core of the book. The coverage of these two chapters is arranged alphabetically, first by family, then by genera, and finally species. The selection of species covered was made to provide examples of virtually all of the native genera and close to half of the native species of Hawaiian plants. Discussion of each family and genus in the flora begins with a brief introductory paragraph describing the key morphological features, geographic distribution, and life-forms of that family or genus. Each species illustrated with a photograph has a brief narrative paragraph describing whether it is endemic or indigenous, its life-form, conservation status, key morphological characteristics, native habitat and elevational range, and occurrence within the Hawaiian Islands. Native Hawaiian names and indigenous uses are included. Although this arrangement generally follows that of the Manual of the Flowering Plants of Hawai‘i, we have chosen to update the taxonomic status of each systematic level to represent the most recent insights on phylogeny and classification gained primarily from molecular studies. Data from DNA sequencing studies over the past decade have produced a new understanding of generic and family relationships among vascular plants and have thus led to changes in the traditional delineations of these systematic units. We have adopted changes in the current acceptance for names at all taxonomic levels made since the publication of the Manual and Palmer (Fern Flora), as shown in the online checklist of the Smithsonian Institution (available at http:// botany.si.edu/pacificislandbiodiversity/hawaiianf lora/index .htm). As examples of these changes, the new delineation of the Malvaceae has been broadened to include the Sterculiaceae, and the Asclepiadaceae has now been subsumed as a subfamily within the Apocynaceae. The former Myrsinaceae is now placed within the Primulaceae, while former members of the Flacourtiaceae are now included in the Salicaceae and the former members of the Chenopodiaceae are now subsumed within the Amaranthaceae. We leave the other changes in family and generic ranks as

xi

xii

P R E FA C E

surprises to those not familiar with the new consensus phylogenies. To minimize confusion, we have cross-indexed the text for these changes in names that differ from those used in the 1990 edition of the Manual and Fern Flora. Following the two systematic chapters of photographs and species descriptions, the book closes with three final chapters. Chapter 6 describes and illustrates the key species brought to the Hawaiian Islands by the Polynesian voyagers. Chapter 7 focuses on the invasive alien species that are widely present in the Hawaiian Islands today. Beyond illustrating the most significant of these invasive alien species, this chapter discusses the ecological impact of these invasive species in altering natural community structure, reducing biodiversity, and altering ecosystem processes. Finally, chapter 8 presents an assessment of the present status of plant conservation in the Hawaiian Islands and suggests future scenarios for the maintenance of habitat protection and biodiversity. An appendix at the end of the book presents a complete checklist of the 1,352 vascular plant taxa considered to be native to seventeen main islands in the Hawaiian Archipelago at the time of this writing. Families are arranged alphabetically, as are the individual genera and species within the dicots, monocots, and fern and fern allies. Specific information is given for each taxon on whether endemic or indigenous, island distribution, their conservation assessment, and federal conservation status. In preparing this book, we thank many people for their help in providing information, slides, and general moral support. In particular, we acknowledge with great appreciation the authors of Manual of the Flowering Plants of Hawai‘i and Hawai‘i’s Ferns and Fern Allies, whose pioneering work formed the basis for our plant descriptions as well as this project in its entirety. Many of the photographs in this book would never have been possible without the assistance of a group of wonderful individuals. We hold these men and women in the highest esteem for their knowledge and invaluable assistance in the field and for allowing use of their photographs. Their individual and collective contributions have been truly remarkable in advancing our understanding of the flora of the Hawaiian Islands. We list these individuals in alphabetical order by island: Kaua‘i—Tim Flynn, David Lorence, Steve Perlman, William Theobald, and Ken Woods; O‘ahu— Gerry Carr, Wayne and Betsy Gagne, Joel Lau, Ken Nagata, John Obata, Clyde Imada, Dan Palmer, Sy Sohmer, and Grady Timmons; Moloka‘i—Joan Aidem; Maui—Robert Hobdy, Angela Kay Kepler, Lloyd Loope, Art Medeiros, and Forest and Kim Starr; Hawai‘i—Mae and William Mull, Lani Stemmerman, and Dietrich Varez; Mainland—Sherwin Carlquist, William Martin, Robert Ornduff, Don Reynolds, Anne Sakai and Steve Weller, Warren Wagner, and Laurel Woodley. For any individuals that we have inadvertently omitted, our thanks go out to you as well.

Note Recent paper publications have resulted in changes affecting accepted names and classifications of the Hawaiian flora. Vernon, A. L., and T. A. Ranker. 2013. Current status of the ferns and lycophytes of the Hawaiian Islands. American Fern Journal 103:59–111. The Hawaiian genera of Woodsiaceae are now split between Athyriaceae (Athyrium, Deparia, and Diplazium) and Cystopteridaceae (Cystopteris). The Hawaiian species of Grammitis are now placed in Oreogrammitis (Polypodiaceae). Grammitis baldwinii to Oreogrammitis baldwinii (Baker) Parris Grammitis forbesiana to Oreogrammitis forbesiana (W. H. Wagner) Parris Grammitis hookeri to Oreogrammitis hookeri (Brack.) Parris—also now considered to be endemic, not indigenous Lu, P. L., and C. W. Morden. 2014. Phylogenetic relationships among dracaenoid genera (Asparagaceae: Nolinoi­ deae) inferred from chloroplast DNA loci. Systematic Botany 39:90–104. Hawaiian species of Pleomele are sister to all other dracaenoids and have now been moved to a new endemic genus, Chrysodracon. Pleomele aurea to Chrysodracon aurea (H. Mann) P.-L. Lu & Morden Pleomele auwahiensis to Chrysodracon auwahiensis (H. St. John) P.-L. Lu & Morden Pleomele fernaldii to Chrysodracon fernaldii (H. St. John) P.-L. Lu & Morden Pleomele forbesii to Chrysodracon forbesii (O. Deg) P.-L. Lu & Morden Pleomele halapepe to Chrysodracon halapepe (H. St. John) P.-L. Lu & Morden Pleomele hawaiiensis to Chrysodracon hawaiiensis (O. Deg. & I. Deg.) P.-L. Lu & Morden

Korthalsella latissima

Lāna‘ihale, Lāna‘i

CHAPTER 1

Landforms and Climate The directional line of the Emperor Seamounts is north to south. About 40 Ma, something happened to rotate the direction of the Pacific Plate movement to its present northwest-to-­ southeast direction in the Hawaiian Islands. It is not clear what caused this rotational shift, but it occurred about the same time as the Indian subcontinent collided with the Eurasian Plate. Kure Atoll, the westernmost of the existing Hawaiian Islands, was formed about 30 Ma. From this atoll, the age of the Northwestern Hawaiian Islands decreases steadily moving southeastward across each island or atoll.

Landforms and Geologic History The distinctive linear shape of the Hawaiian Island chain has been produced through geologic time as the Pacific Plate has slowly moved across a deep hot spot in the Earth’s mantle. Heat from the hot spot produces a continuous source of magma by melting the overriding Pacific Plate. The magma rises through the mantle and crust to erupt onto the seafloor, gradually building a shield volcano to a height thousands of meters above the deep ocean basin, until it finally emerges above sea level. The Hawaiian hot spot is located today beneath the southeastern portion of the Big Island of Hawai‘i. This process produces a cycle of island birth and death as the Pacific Plate continues to move across the hot spot at a rate of 5–10 centimeters (cm) per year. When the movement of the plate carries an island beyond the hot spot, removing it from its source of magma, the volcano becomes extinct. As one island volcano becomes extinct and begins to slowly erode from heavy rain and massive landslides, another island is developing over the hot spot, and the cycle continues. This cycle can be seen in the Northwestern Hawaiian Islands, all once volcanic islands that have largely disappeared. In the tropics, coral begins to grow around an island, forming a fringing reef. As the island erodes and sinks, a lagoon separates the island from a barrier reef. When erosion finally removes the portions of the landmass above sea level, an atoll is formed, leaving a coral reef circling the top of the submerged volcanic platform. When the rate of erosion and subsidence becomes greater than the growth rate of the coral reefs, the coral eventually dies and the atoll sinks below the surface to become a flat-topped seamount. The Hawaiian chain actually begins not with the Northwestern Hawaiian Islands but rather with the Emperor Seamounts, which extend from near the Aleutian Trench at the margin of the Pacific Plate south in a line to the present Kure Atoll. These seamounts represent a chain cycle of volcanic island formation over the Pacific Plate hot spot that includes at least 129 volcanoes. The oldest seamount, near the Aleutian Trench, was once an island formed about 75–80 million years ago (Ma), and seamounts to the south become increasingly younger in age of formation. The Kōkō Seamount, located today at the southern end of the Emperor Seamounts, was the last large island in this chain and submerged about 33 Ma, leading to a period of several million years in which no islands existed.

The Northwestern Hawaiian Islands The Northwestern Hawaiian Islands are a chain of islands that extend along a linear path approximately 1,600 kilometers (km) southeast from Kure Atoll to Nīhoa Island to the west of Kaua‘i. These islands, which with one exception are administratively part of the State of Hawai‘i, are remnants of once larger islands that have slowly eroded and subsided. Today they exist as small landmasses or coral atolls that cover the remnants of the volcanic islands. Although these islands have been given Hawaiian names in modern times, with the exception of Nīhoa these are not names that were used prior to the arrival of Euro-Americans to the Hawaiian Islands. Kure Atoll (Hawaiian: Kānemiloha‘i) is the westernmost of the Hawaiian chain, lying about 160 km east of the international date line. Kure has the distinction of being the northernmost coral atoll in the world. The volcanic landmass age of the island is about 30 Ma. It consists of a 10 km-wide, nearly circular barrier reef surrounding a shallow lagoon and small sand islets, with a total land area of only 86 hectares (ha) (0.86 km2) and a maximum elevation of 6 m above sea level. The only land of significant size is called Green Island and is habitat for hundreds of thousands of seabirds. Kure lies close to the latitude at which reef growth just equals reef destruction by various physical forces. As Kure continues to be slowly carried along to the northwest by the motion of the Pacific Plate, it is expected to move into waters too cold for coral and coralline algae growth to keep up with isostatic subsidence of its volcanic base. At this stage, it will gradually become a submerged seamount. The atoll is named for Captain Kure, a Russian navigator who is reputed to have discovered the atoll in the early nineteenth century. Midway Atoll (Hawaiian: Pihemanu), whose name indicates its approximate position halfway between the North American

1

2

LANDFORMS AND CLIMATE

1.1 

Hawaiian Island archipelago

and Asian continents, is an atoll located about 90 km east of Kure. It consists of a ring-shaped barrier reef and several sand islets, with 6.4 km² of land area and a maximum elevation of 4 m. This size makes it the largest of the Northwestern Hawaiian Islands and the most well known because of its past history as an

1.2 

Kure Atoll

airport midway between North America and East Asia. The volcanic landmass age of the island is 28.7 Ma. The two significant areas of land, Sand Island and Eastern Island, provide important habitat for seabirds but have been highly impacted by human activities in the past. Except for Midway, all of the Northwestern Hawaiian Islands are administratively part of the State of Hawai‘i. Midway is considered an unincorporated territory and is administered by the U.S. Department of the Interior. Pearl and Hermes Atoll (Hawaiian: Holoikauaua) consists of a few small sandy islands contained within a lagoon and surrounded by a coral reef. The reef is virtually devoid of vegetation except for small patches in wet years. Pearl and Hermes Reef has a volcanic landmass age of 26.8 Ma. The total land area of all its islets is only 36 ha (0.36 km2), with a maximum elevation of 3 m. In addition, the atoll here includes 404 km2 of submerged reefs. The atoll bears the name of two English whaling ships that were wrecked on the previously unknown reef in 1822. The endangered Laysan finch has been introduced to the island to provide a backup population should a disaster wipe out the endemic population on the island of Laysan. Lisianski (Hawaiian: Papa‘āpoho) is a small volcanic island with a land area of about 1.5 km² and a maximum elevation of about 12 m. The volcanic landmass age of the island is 23.4 Ma. A sand-filled depression located between two tall dune areas on the island is thought to once have been a lagoon like the one on Laysan, its nearest neighbor. Linked to Lisianski are the extensive Neva Shoals, a reef that covers a broad area of 979 km². The island was named after its discoverer, Captain Urey Lisiansky of the Russian ship Neva, which went aground on the reef in 1805.

LANDFORMS AND CLIMATE

1.3 

Laysan Atoll

Laysan Island (Hawaiian: Kauō) comprises a landmass located about 1,500 km west of Honolulu and extending about 1.6 by 2.4 km in size, with a land area of 4.1 km² and a maximum elevation of 12 m. It is an atoll, although the land completely surrounds a shallow central lake lying 2.4 m above sea level. Lake Laysan, one of only five natural lakes in the Hawaiian Islands, is hypersaline, with a salt concentration three times that of the ocean. The volcanic landmass age of the island is 20.7 Ma. The island’s natural ecosystems, rich in endemic bird taxa and including undescribed species of Pritchardia and sandalwood (Santalum ellipticum), were largely destroyed by human influence around the turn of the twentieth century when guano harvesting and the commercial introduction of rabbits eliminated most of the vegetation. At least 26 plant species have been extirpated, and 3 endemic bird taxa (Laysan ‘apapane, Laysan rail, and Laysan millerbird) once found on the island are now extinct. The endemic Laysan finch and Laysan duck are endangered today. Pollen cores from the central lagoon of Laysan contain evidence of coconut palm, suggesting that there might have been an early colonization of the island by Polynesians. The island name was bestowed by American whaling ships in the early part of the nineteenth century. Maro Reef (Hawaiian: Nalukakala) is a largely submerged coral atoll with about 1.4 ha of semidry land, lying 1,370 km west of Honolulu. This small area can be submerged with high tides. Maro Reef forms the largest reef area in the Northwestern Hawaiian Islands. Gardner Pinnacles (Hawaiian: Pūhāhonu), located 946 km northwest of Honolulu, form the westernmost exposed volcanic remnants in the Hawaiian archipelago. The pinnacles consist of two barren volcanic outcrops surrounded by a reef. The total area of the two small islets, remnants of an ancient volcano, is about 2.4 ha (0.024 km2). The tallest peak reaches a height of 58 m. The age of these two volcanic remnants is 15.8 Ma. The surrounding reef area is very large. The island was named in honor of Edmund

Gardner, captain of one of the first American whaling ships to reach the Hawaiian Islands in 1819. French Frigate Shoals (Hawaiian: Mokupāpapa) is the easternmost atoll in the Hawaiian Archipelago. The circular atoll is 29 km across, with many small sandy islands around its fringes. The total land area is about 25 ha, and the islets are mostly no more than 4 m in elevation. The small size was sufficient to have supported a U.S. Navy landing field in World War II. The volcanic landmass age of the island is 12.8 Ma. The shoals have an associated coral reef area of 938 km². La Pérouse Pinnacle, a tiny basalt remnant of the volcanic origins of French Frigate Shoals, is 37 m tall. The name of the shoals came from their discovery by

1.4 

Nīhoa Island

3

4

LANDFORMS AND CLIMATE

French captain Jean François La Pérouse, who narrowly missed running aground here in 1786. Necker Island (Hawaiian: Mokumanamana) is a rocky volcanic island 249 km northwest of Nīhoa and 690 km northwest of Honolulu. It has a land area of only 18 ha (0.18 km2) and a maximum elevation of 84 m. The landmass age of the island is 11.0 Ma. The name of the island was given by Captain La Pérouse in honor of the French minister of finance. Nīhoa is a rocky volcanic island located 450 km northwest of Honolulu, with a land area of 70 ha (0.70 km2) surrounded by 570 km² of coral reef. Its steep topography and crater shape show its volcanic origin. Nīhoa is the highest of the Northwestern Hawaiian Islands, with two areas of moderate elevation—Miller’s Peak (275 m elevation) in the west and Tanager Peak (260 m elevation) in the east. The landmass age of the island is 7.3 Ma. Despite its size, the island has sufficient soil to have supported a small Hawaiian population from about AD 1000 to 1700. Two endemic bird taxa, the Nīhoa finch and Nīhoa millerbird, are endangered. The palm Pritchardia remota is endemic to this island and Ni‘ihau. On 15 June 2006, President George W. Bush signed a bill designating the Northwestern Hawaiian Islands Marine National Monument, with the name changing the following years to Papahānaumokuākea Marine National Monument. The monument encompasses approximately 362,600 km2 and includes 11,650 km2 of coral reef habitat. Its creation consolidates the previously existing Hawaiian Island Reservation established by Theodore Roosevelt in 1909, the Hawaiian Islands National Wildlife Refuge, and the Midway Atoll National Wildlife Refuge.

1.5 

Hawaiian Islands

The Major Islands The Hawaiian Islands include eight major islands: Ni‘ihau, Kaua‘i, O‘ahu, Moloka‘i, Maui, Lāna‘i, Kaho‘olawe, and Hawai‘i.

NI‘IHAU Ni‘ihau is a small, elongate island approximately 29 km in length by 10 km in width and about 180 km2 in area. The maximum elevation (Pānī‘au) reaches only 381 m. Ni‘ihau comprises the low-lying remains of a volcanic shield built about 5.2 Ma. The summit of the volcano was originally northeast of the present-day island but was subsequently removed, along with much of the northeast side of the island, by erosion and a massive landslide. Human disturbance, primarily ranching, has drastically changed the vegetation and hydrological parameters of Ni‘ihau, leaving only small native vegetation communities. Ni‘ihau has been privately owned since 1864, and access has been and continues to be restricted. Ka‘ula is a small, crescent-shaped island lying about 32 km west-southwest of Ni‘ihau, with an area of 64 ha (0.64 km2). The island is the upper portion of a cone of volcanic tuff that rests on top of a larger, submerged shield volcano. At its highest point, the island reaches an elevation of 167 m. It is slightly older than Ni‘ihau, with a landmass age of 5.3 Ma. Lehua is another small island lying just north of Ni‘ihau, with an area 1.15 km2 and a maximum elevation of 213 m. This barren island is a tuff cone that was once part of the now extinct Ni‘ihau volcano.

LANDFORMS AND CLIMATE

Kaua‘i is the fourth largest of the main Hawaiian Islands, with an area of 1,430 km². The island formed about 4.7 Ma as a single shield volcano. Subsequent collapse of portions of the caldera occurred, followed by a second period of volcanic flows that filled the caldera and produced the flat-lying Alaka‘i Plateau. More than 1.5 million years after the primary shield-building stage had ceased on Kaua‘i, renewed volcanism produced new flows over two-thirds of the eastern side of the island. The highest point on Kaua‘i is Kawaikini at 1,598 m, followed by Mount Wai‘ale‘ale near the center of the island at 1,569 m. The windward upper slope of Mount Wai‘ale‘ale is one

of the wettest spots on earth, with a mean annual rainfall of 11,700 millimeters (mm) and a record 17,340 mm of rainfall in 1982. The runoff from this high annual rainfall has eroded deep valleys on the island, carving out canyons with many hanging waterfalls. The largest of these is Waimea Canyon, which drains the Alaka‘i Plateau. It measures 16 km in length, 3 km in width, and more than 1,000 m in depth. The Kalalau Lookout above the Nāpali Coast on the northwest shore of Kaua‘i has an elevation of 1,256 m. Due to its age and relative isolation, Kaua‘i is second among the islands for the highest levels of floristic diversity, and it boasts the highest endemism in the Hawaiian Archipelago (chapter 2).

1.6 

1.7 

KAUA‘I

Waimea Canyon, Kaua‘i

1.8 Wai‘ale‘ale, Kaua‘i

Kalalau Lookout, Kaua‘i

5

6

LANDFORMS AND CLIMATE

1.9 Wai‘anae Mountains, O‘ahu

1.10 

Summit Ridge, Ko‘olau Mountains, O‘ahu

1.11 

Waterfall, Castle Trail, leeward Ko‘olau Mountains, O‘ahu

However, the vegetation on Kaua‘i has undergone extreme alterations, as with the other islands, first by Native Hawaiians and more recently with land conversion to agricultural and pasture use by Euro- and Asian-Americans. Native forests are now largely restricted to mesic and wet areas at higher elevations.

O‘AHU O‘ahu, the third largest Hawaiian island, extends about 71 km in length and 48 km in width, with a total area of 1,545 km². The island is formed from the remnants of two large shield volcanoes, the older Wai‘anae volcano to the west and the younger Ko‘olau volcano to the east. Their original shield volcano shape has been lost as a result of extensive erosion and massive subaerial landslides, and today these volcanoes are mountain ranges consisting of long, narrow ridges. The Wai‘anae Mountains, 35 km in length, with narrow ridges and steep slopes as the predominant features, were formed about 3.0 Ma along a northwest to south-southeast rift zone. The caldera lies between the north side of Mākaha Valley and the head of Nānākuli Valley. The striking features of the range are the great amphitheater-shaped valleys that slope up to the steep pali at the back of the valleys. Ka‘ala, the highest point in the range as well as on the island, reaches an elevation of 1,220 m. The summit of Ka‘ala is a plateau formed by thick ‘a‘ā lava flows, with poor drainage conditions that produce a montane bog. Pu‘u Kalena near Ka‘ala is the second highest peak on O‘ahu at 1,068 m. The Ko‘olau Mountains were built by eruptions that likewise took place along a northwest-trending rift zone, forming a range approximately 60 km in length. The Ko‘olau Mountains today represent the western half of the original volcano. The eastern half of the volcano, including much of the summit caldera, was lost to a cataclysmic landslide into the Pacific Ocean in past geologic time. These remains exist today as massive fragments strewn over 160 km of the ocean floor to the northeast of O‘ahu.

LANDFORMS AND CLIMATE

1.12 

Koko Crater, O‘ahu

The Ko‘olau volcano attained its maximum elevation about 2.5 Ma, when it is thought to have reached as high as 3,000 m. Subsequent erosion, subsidence, and the major underwater landslide of its eastern half have since greatly reduced its size, but steep slopes remain. The highest point today is Pu‘u Konahuanui at 960 m. Scattered eruptions over the past 500,000 years on the flanks of Ko‘olau volcano have created many of the landmarks around eastern O‘ahu, such as Diamond Head (232 m), Hanauma Bay, Koko Head (196 m), Punchbowl Crater (152 m), Tantalus (614 m), and Āliapa‘akai (Salt Lake).

MAUI NUI COMPLEX The Maui Nui complex of the Hawaiian Islands consists of the islands of Moloka‘i, Maui, Lāna‘i, and Kaho‘olawe, which were connected in the past as a single landmass. Another small volcano west of Moloka‘i, known as Penguin Bank, is now completely submerged and covered with a cap of coral reef. Interestingly, this bank reached its maximum shield volcano stage about 2.2 Ma and at that time was connected by a land bridge to O‘ahu, which was further connected to West Moloka‘i, well before the development of Haleakalā. The connections from O‘ahu to Penguin Bank and to West Moloka‘i became submerged about 2.0 and 1.9 Ma, respectively, and thus the Maui Nui connection to O‘ahu lasted for only about 0.3 Ma. The seafloor between the four Maui Nui islands is relatively shallow, about 500 m deep, in comparison to the outer edges of the combined island mass, where the ocean floor plummets

to the abyssal ocean floor. At its greatest extent around 1.2 Ma, Maui Nui was larger than the modern island of Hawai‘i, but the individual islands became separated during subsequent subsidence beginning around 0.6 Ma. However, low sea levels during interglacial periods allowed the Maui Nui complex to form again for brief periods. Overall then, the Maui Nui complex has formed a single large landmass for more than 75% of its geologic history.

MOLOKA‘I The island of Moloka‘i, the fifth largest in the Hawaiian Islands chain, is approximately 61 km in length and up to 17 km in width, with an area of about 673 km2. Two shield volcanoes, West Moloka‘i and East Moloka‘i (Wailau), make up most of the landmass of the island, first forming volcanic shields about 2.0 and 1.8 Ma, respectively. Newly active volcanic flows about 300,000 years ago formed Kalaupapa Peninsula on the northern face of the island. West Moloka‘i lacks high elevations, reaching only 436 m at Mauna Loa. This low western half of the island is dry, and the soil is heavily denuded due to heavy grazing under past land management practices. The Mo‘omomi Dunes on the northwest coast provide one of the few remaining areas of intact coastal shrublands in the Hawaiian Islands. East Moloka‘i, the larger volcano forming two-thirds of the island, reaches an elevation of 1,512 m on Kamakou peak. Topographically, the windward side of East Moloka‘i differs from the leeward south side, with steep cliffs along the northern coast and

7

8

LANDFORMS AND CLIMATE

1.13 

East Moloka‘i

deep inaccessible valleys dissecting the coastal area. These steep cliffs, the highest sea cliffs in the world, are the result of a massive underwater landslide that removed most of the northern half of East Moloka‘i about 1.4 Ma. The soils of the upper crest of the volcano are poorly drained, producing bogs high in organic matter. The Pēpē‘ōpae Bog lies near the summit of Kamakou. The gulches and valleys are usually very steep, but a few are gently sloping. Much of the native vegetation on the northern part of East Moloka‘i is relatively intact by Hawaiian standards because of its general inaccessibility to humans and nonnative animals.

of central Lāna‘i is relatively flat, several deep canyons have been eroded into the slopes of the old volcano. The largest of these is Maunalei Gulch, which provides most of the drinking water for the island. Tall sea cliffs on the southwest shore of Lāna‘i reach 330 m in height. As with other areas of the Islands, heavy grazing of domestic and feral animals in the nineteenth century destroyed much of the native vegetation, eroded soils, and caused major deforestation. Much of the vegetation has never fully recovered, although there have been serious attempts to revegetate upland areas, mostly with introduced species.

LĀNA‘I

KAHO‘OLAWE

Lāna‘i is a relatively small, kidney-shaped island with a total area of 364 km2. It was formed by a single shield volcano that developed about 1.5 Ma along three rift zones. As part of Maui Nui, the Lāna‘i volcano grew on the flank of the older East Moloka‘i volcano and was later partly buried by the younger West Maui volcano. The highest point on the island is Lāna‘ihale, which reaches 1,026 m in elevation. The former summit caldera is largely filled with hardened lava today. Although the upland area

Kaho‘olawe is the smallest of the major Hawaiian Islands, with an area of only 116 km² and dimensions of about 18 by 11 km. The island was formed by a single shield volcano about 1.2 Ma. The topography consists of a nearly filled caldera and a rift zone that trends to the southwest, with the highest point being 452 m at the crater of Lua Makika at the summit of Pu‘u Moa‘ulanui. The island is relatively dry because of its low elevation and its position in the rain shadow of Haleakalā.

1.14 Lāna‘ihale, Lāna‘i

1.15 

Eroded landscape of Kaho‘olawe

10

LANDFORMS AND CLIMATE

Kaho‘olawe has had a complicated history of human exploitation that has affected its terrain and environment, leaving it dry and barren. Tens of thousands of sheep and goats grazed on the island in the nineteenth and early twentieth centuries. Resulting overgrazing destroyed most of the vegetation of the island, with subsequent erosion removing much of the topsoil. More than one-quarter of the island has been eroded down to saprolitic hardpan. In 1939, the United States military commandeered the island for use as a bombing and artillery target, and military control continued until 1993 when the U.S. Congress returned Kaho‘olawe to the State of Hawai‘i.

MAUI Maui, the second largest of the Hawaiian Islands, has an area of 1,884 km². The island is formed by two large volcanoes separated by a low-lying isthmus. The older West Maui volcano completed its shield-forming stage of development about 1.5 Ma. After about 500,000 years of dormancy, smaller eruptions from cinder cones occurred about 380,000–600,000 years ago. Stream erosion has subsequently cut deep valleys and ridges into the originally shield-shaped volcano, which channel upward to the peak of Mount ‘Eke at 1,764 m. The West Maui volcano is considered extinct today. Much of the runoff of the high rainfall from the West Maui Mountains flows through the ‘Īao Valley, where the ‘Īao Needle rises 370 m above the valley floor.

1.16 Mount ‘Eke, West Maui

The larger and younger Haleakalā volcano of East Maui reaches an elevation of 3,055 m. Below the summit lies a massive crater approximately 11 km across, 3 km wide, and nearly 800 m deep. The interior walls are generally steep and the crater barren-looking, with a scattering of volcanic cinder cones. The crater rim is irregular in height and dips to 2,500 m at Kaupō Gap, allowing for the entrance of dense clouds and extensive plant cover. Haleakalā’s crater is not technically a caldera but instead was formed when the headwalls of two large erosional valleys merged at the summit of the volcano. These valleys formed the large gaps of Ko‘olau on the north side and Kaupō on the southeast of the Haleakalā depression. The volcano’s shield stage of development ended about 1.2 Ma, when it probably reached an elevation of about 3,500 m. Haleakalā is considered a dormant volcano, as there have been numerous small eruptions in the last 30,000 years along two rift zones on its southwestern and eastern flanks. The last of these may have occurred about 1790 at La Pérouse Bay. The isthmus joining West and East Maui was created during the shield-building stage of Haleakalā as lava flowed into West Maui. It is bounded by two embayments, one to the north and one to the south. Molokini is a partially submerged volcanic crater lying between Maui and Kaho‘olawe, both part of Maui County in Hawai‘i. It has an area of 9 ha and reaches a height of 49 m above sea level.

1.17 

‘Īao Valley, West Maui

1.19 

East Maui watershed

1.18 

Haleakalā, East Maui

12

LANDFORMS AND CLIMATE

HAWAI‘I Hawai‘i, the Big Island, is the largest of the Hawaiian Islands by far, with its total area of 10,433 km2 comprising almost twothirds of the total land area of the entire State of Hawai‘i. The island was formed by five large shield volcanoes that erupted somewhat sequentially, thus allowing younger flows to cover older flows. The oldest of these volcanoes is the extinct Kohala Volcano on the northern margin of the island, which reaches 1,670 m in elevation. The volcano emerged above sea level more than 600,000 years ago. At the end of its shield-building stage 250,000 to 300,000 years ago, a massive landslide removed the volcano’s northeast flank, leaving steep cliffs that drop to the coast. This landslide cut back to the summit of the volcano, which at the time was more than 1,000 m higher than today. The Kohala Volcano is considered to be in a transition between the post-shield and erosional stages of its life cycle. The broad shield of Hualālai forms the western margin of Hawai‘i and reaches an elevation of 2,521 m. It is estimated that the volcano reached its shield stage of development about 400,000 years ago. The oldest dated rocks on the surface have an age of about 128,000 years. The west-facing flank of Hualālai

1.20 

Kohala Mountains, Hawai‘i

forms a large underwater slump. A major feature of Hualālai is Pu‘u Wa‘awa‘a, a volcanic cone rising 372 m in height and measuring over 1.6  km in diameter. Much of the surface slopes of Hualālai consist of lava flows less than 5,000 years in age. Although dormant today, six vents opened enough to produce lava flows over a period from the late 1700s to 1801, with two of these generating lava flows that reached the sea along the west coast of the island. Next in age is Mauna Kea, the highest point on the island, reaching 4,205 m in elevation. Measured from its true base in the deep ocean basin some 6,000 m below sea level, Mauna Kea is the tallest mountain in the world, well surpassing Mount Everest. The early eruptions of Mauna Kea began about 1.0 Ma, and the volcano reached its maximum shield stage about 200,000–250,000 years ago. Small local eruptions 4,000–6,000 years ago formed numerous overlapping cinder cones that filled the caldera. These cinder cones now form the peaks at the summit. While there have been no historical eruptions, Mauna Kea is considered to be a dormant rather than an extinct volcano. The summit area of the volcano was glaciated four times during the pluvial periods of the Pleistocene, with a glacial area 150 km2 extending down to an elevation of 3,300 m at its maximum extent.

1.21 

Hāmākua Coast, Hawai‘i

1.22 

Mauna Kea, Hawai‘i

14

LANDFORMS AND CLIMATE

1.23 

Mauna Loa summit, Hawai‘i

The last melting of glaciers occurred about 11,000 years ago. Snowfall occurs irregularly today in winter at elevations above about 3,400 m. Mauna Loa is a massive shield volcano that continues to be active today. Its large summit caldera, 3–5 km across, reaches 4,169 m, just below the elevation of Mauna Kea. Despite this elevation today, there is no evidence of any glacial activity on the summit during the Pleistocene. This massive volcano reached sea level somewhere about 0.6–1.0 Ma, and it has continued to grow, although more slowly, over the last 100,000 years. Nearly 98% of the surface area of Mauna Loa is covered by lava flows less than 10,000 years in age. Eruptions today generally occur at the summit and in two rift zones that extend northeast and southwest of the summit. The caldera is thought to have formed only about 1,000–1,500 years ago when a large eruption from the northeast rift zone emptied the magma chamber and collapsed the summit. The highly fluid lava flowing in the past from the summit of Mauna Loa has given it shallow slopes in comparison to Mauna Kea. Lava flows from Mauna Loa have overlapped older flows from Mauna Kea and Hualālai to form a broad saddle area lying at about 1,500–2,200 m elevation at the convergence of the three volcanoes at the center of the island. Kīlauea, the youngest of the shield volcanoes on the Big Island, lies against the southeast flank of much larger Mauna Loa.

It reaches an elevation of 1,248 m, with a summit caldera about 165 m deep and measuring about 3 by 5 km in size. The earliest formation of the Kīlauea Volcano began about 300,000–600,000 years ago and reached above sea level about 50,000–100,000 years ago. However, the oldest dated surface flows are only 23,000 years old. About 90% of the surface of Kīlauea is covered by lava flows less than 1,100 years old, and 70% of the surface is thought to have flows younger than 600 years. Historical eruptions of Kīlauea have occurred primarily from the summit caldera and along the East and Southwest rift zones. Eruptions have been almost continuous since 1983, with many of the lava flows reaching to the sea. Lava flows in 1987 buried much of the Royal Gardens subdivision, near the boundary of Hawai‘i Volcanoes National Park. Three years later, new flows destroyed the towns of Kalapana and Kaimū as well as Kaimū Bay, the Kalapana black sand beach, and a large section of Route 130 which now dead-ends at the lava flow. The last remaining home in the Royal Gardens subdivision was destroyed by a flow in 2012. Lō‘ihi is an actively erupting seamount that lies 975 m below sea level to the southeast of Hawai‘i. It is probable that continued volcanic activity from Lō‘ihi will cause the volcano to eventually reach sea level and later attach at the surface onto Kīlauea, adding even more land to Hawai‘i. This is not expected to occur for tens of thousands of years.

1.24 

Mauna Loa slope, Hawai‘i

1.26 Ka‘ū District, Hawai‘i

1.28 

Kīlauea lava flow at Kalapana, July 2010

1.25 

Kīlauea Iki, Hawai‘i

1.27 

Eruption between Pu‘u Ō‘ō and Nāpau Crater, Hawai‘i, March 2011

16

LANDFORMS AND CLIMATE

Climate Lying just inside the Tropic of Cancer between latitudes 19 and 22 degrees north and surrounded by water, the Hawaiian Islands experience a moderated tropical environment. Daytime high temperatures in the lowlands seldom exceed 32 degrees centigrade (90 degrees Fahrenheit), while extreme low temperatures at night are above 10 degrees centigrade (50 degrees Fahrenheit). Differences of mean daily temperatures in summer and winter are less than 4 degrees centigrade (39 degrees Fahrenheit). This is smaller than the amplitude of mean temperature change from day to night. Low temperatures do occur on the high volcanoes, where frosts are common. The low temperature record for the Islands is –13 degrees centigrade (–9 degrees Fahrenheit) on the summit of Mauna Kea.

PATTERNS OF RAINFALL The topographic form and orientation of the landmasses of the individual Hawaiian Islands have a major effect on rainfall patterns. This influence can be seen by considering that the typical levels of annual rainfall are 500–750 mm over the open ocean around the islands. On the islands themselves, however, local conditions produce heavy rainfall of more than fifteen times this amount on mountain peaks and as little as a third of this amount on leeward slopes in the rain shadow of the higher volcanoes. In addition, multiyear cycles of ocean-atmosphere interactions such as El Niño–Southern Oscillation (ENSO) events and cycles of the Pacific Decadal Oscillation (PDO) can significantly influence amounts of rainfall. The highest amounts of annual rainfall in the Hawaiian Islands are associated with orographic effects acting on the northeastern trade winds. The high landmasses of the islands force the moist air mass of the trade winds upward, cooling them and producing heavy amounts of rainfall at the windward crest of mountain ridges. These rains often begin in late afternoon and continue overnight, ceasing again in the morning. The peak of this rainfall typically occurs at an elevation of 1,500–1,800 m, where ridgelines are often obscured by cloud cover. An exception to this pattern of high rainfall on the windward slopes at this elevation occurs when the windward face of the mountains is extremely steep, as with the Ko‘olau Mountains of O‘ahu and the windward face of East Moloka‘i. Under these topographic conditions, the rapid upward movement of the moist trade winds produces high winds at the crest of the ridges that push the peak amount of rainfall across to the upper leeward face. Above 2,000 m elevation on the higher volcanoes, an inversion of warm air limits the upward movement of the trade winds, and rainfall decreases significantly. Thus, the summits of the high volcanoes on Hawai‘i are arid, with less than 400 mm of annual precipitation. Summer is the drier season for most of the Hawaiian Islands. This is strongly true of the more arid leeward sides of the islands and less strongly so on the upper windward slopes of the windward sides, which are influenced by the strong dominance of the trade winds even in summer. The one exception to this pattern occurs with the special conditions on the leeward Kona Coast of Hawai‘i, where more summer rainfall occurs. Well more than half of the rainfall on the drier leeward sides of the islands comes from a small number of winter storms. These

include rains occurring with the passage of cold fronts and Kona storms. Kona storms are associated with a change in the prevailing wind pattern, with south and southwesterly winds replacing the trade winds. These storms are the result of upper-level low-pressure systems that become “cut off” from the main flow of the middle-latitude westerly wind belt. Once formed, these lows may remain relatively fixed in position over the islands for days by the blocking action of high-pressure systems. Kona storms, which may last for a week or more, are characterized by intense rains interrupted at times by intervals of lighter rain or even partial clearing. Remarkably heavy rainfall can occur under these conditions. A storm in January 1957 dropped more than 965 mm of rain in a twenty-four-hour period on Kīlauea Sugar Plantation, Kaua‘i, with 150 mm falling in one thirtyminute period. Typically there are two to five Kona storms each winter, plus a similar number of cold front storms. Hurricanes are relatively uncommon in Hawai‘i, with five occurrences over the past sixty years. These have often been associated with years with El Niño–Southern Oscillation conditions, as with Hurricane ‘Iniki in 1992, which caused an estimated $2.3 billion in storm damage. In four of these hurricanes, Kaua‘i has received the major damage from the storms. Less intense tropical storms pass close enough to the Islands to impact weather conditions every year or two. Unlike cold fronts and Kona storms, which are winter events, hurricanes and less intense tropical storms typically occur in the second half of the year, from July through December.

Rainfall on Individual Islands Lacking sufficient elevations to impact the moist trade winds, rainfall on the low-lying Northwestern Hawaiian Islands is only a small amount higher than precipitation present in the surrounding open ocean. Estimated rainfall levels are 600–750 mm annually. Ni‘ihau is relatively arid because it lies in the rain shadow of Kaua‘i and lacks the elevation needed to catch significant amounts of the moisture of the trade winds. Typical annual amounts of rainfall are only about 300 mm. The rainfall pattern of Kaua‘i is characterized by a maximum level on Mount Wai‘ale‘ale, near the island’s highest point. Steep valleys on both the windward and leeward slopes of the island channel trade winds and Kona storms upslope to the crest of the mountain in all seasons of the year. As a result, Wai‘ale‘ale receives more rainfall than any other instrumented site in the world. The mean annual rainfall here, as already noted, is 11,700 mm. Kekaha, on the leeward west coast of Kaua‘i, receives less than 500 mm of rainfall, despite lying only 25 km southwest of Mount Wai‘ale‘ale. In the Ko‘olau Mountains of O‘ahu, moisture-rich trade winds strike the northeastern face, producing strong uplift with high amounts of rainfall along the crest of the range. Levels of mean annual rainfall are more than 7,000 mm at Kahana, just to the lee of the highest elevation on the windward slopes, and about 4,000 mm at Tantalus above Mānoa Valley. Lower slopes on the leeward side of the range receive about 1,000 mm. Despite lying perpendicular to the direction of the trade winds, the Wai‘anae Mountains lie in the rain shadow of the Ko‘olau Mountains and receive lower amounts of precipitation. Rainfall is high only on Ka‘ala, which receives approximately 2,500 mm annually. This amount would be far greater were it

LANDFORMS AND CLIMATE

not for the rain shadow impact of the Ko‘olau Mountains. Other areas of the Wai‘anae Mountains receive from about 1,500 mm to less than 200 mm in the dry northwestern coastal areas. Levels of rainfall on Moloka‘i are strongly determined by not only topography but the island orientation as well. The highest levels occur on East Moloka‘i, where the orographic uplift of the trade winds produces levels of annual rainfall above 4,000 mm. This amount, while high, is lower than might be expected compared to similar elevations on the windward slopes of other islands. The somewhat lower levels occur primarily because of the east-west orientation of the island, so that its exposure to the trade winds is not as pronounced as with the Ko‘olau Mountains on O‘ahu. The lower elevations of West Moloka‘i are relatively dry, with typical rainfall levels about 750 mm but reaching below 400 mm in places. Rainfall levels are low on both Lāna‘i and Kaho‘olawe because of the rain shadow effect of West Maui. The small area and regular topography of Lāna‘i produce a simple pattern of rainfall. Much of the lowland area of the island receives only 250– 500 mm of annual rainfall, while the higher areas of Lāna‘ihale have 750–1,270 mm of rainfall. Particulate moisture from clouds covering the summit area condenses onto foliage, producing a fog drip that greatly increases the soil moisture available for plant growth. Low topography as well as the rain shadow effect add to the aridity of Kaho‘olawe, where rainfall is very low. Rainfall patterns on Maui are strongly influenced by the orographic effects of the volcanic peaks of West and East Maui in capturing and concentrating trade wind moisture. Mean annual rainfall exceeds 9,000 mm on the summit of West Maui, where deeply incised valleys around the eroded cone funnel air toward the summit from both the trade winds and winter Kona storms, while Big Bog on the windward slope of Haleakalā receives more than 10,000 mm of rainfall. These amounts are second only to Mount Wai‘ale‘ale on Kaua‘i. The other point of maximum rainfall occurs on the northeastern slope of Haleakalā above Hāna. The trade wind inversion concentrates rainfall here at about 650 m elevation, with more than 7,500 mm falling annually. The driest areas of Maui are those lying on the leeward side of the island from Mākena to Kā‘anapali in the rain shadow of the two volcanoes. Lahaina receives an average of only 355 mm annual rainfall. Also moderately dry is the summit of Haleakalā, above the trade wind inversion, which receives about 1,000 mm of annual precipitation, with occasional light snowfall in winter. The varied topography of Hawai‘i produces a complex pattern of rainfall distribution. There are four areas where distinct high levels of rainfall occur. The highest of these are present on the windward slopes of the Kohala Mountains and the eastern slopes of Mauna Loa above Hilo, both associated with orographic effects on trade wind moisture. The Kohala Mountains have their highest rainfall of more than 4,000 mm on the windward slopes of this range near the summit at about 1,600 m elevation. The peak amount of rainfall on Hawai‘i is present at about 760 m elevation on the eastern slopes of Mauna Loa, where

1.29 

Clouds over Haleakalā Crater, East Maui

a strong temperature inversion on the higher volcano limits the upslope flow of moist winds. Here mean annual rainfall exceeds 7,000 mm. A portion of Ka‘ū District, on Mauna Loa’s southeast flank, receives more than 3,000 mm of annual rainfall, an unexpected amount for a slope parallel with the direction of the trade winds. Orographic uplift in this area is enhanced by circulation patterns influenced by thermal heating of the mountain slopes. The final area of high rainfall is similarly surprising. This is the Kona area upslope at 650–900 m elevation from the Kona Coast, which receives an annual rainfall of 1,200–2,500 mm despite its position in the rain shadow of Hualālai and Mauna Loa. Rainfall here results from thermal heating of the large landmasses of the two sheltering volcanoes. This diurnal heating causes air masses to rise, drawing in moisture-laden air from the ocean to the west of Kona, with resulting convergence and light afternoon rains. Regular afternoon cloud cover associated with this convergence produces the conditions for the growth of the famed Kona coffee. Several areas of Hawai‘i are notable for low rainfall and aridity. Kawaihae, which lies in the rain shadow of the Kohala Mountains and Mauna Kea, and the summit area of Mauna Kea itself, both receive only about 200 mm of annual rainfall. Another area of low rainfall is the saddle area lying between Hualālai, Mauna Loa, and Mauna Kea at elevations above the trade wind inversion. This area receives less than 500 mm of rainfall. Similar low levels of rainfall are present at South Point on the southern tip of Hawai‘i, which lies in the lee of Kīlauea. Also appearing very arid is the Ka‘ū Desert, located on the southeastern side of Hawai‘i in the volcano’s rain shadow. Despite its arid appearance, however, this area receives up to 1,200 mm of annual rainfall.

17

Fleshy fruit of Phyllostegia grandiflora

CHAPTER 2

Plant Diversity and Evolution O

with 520 species. Moloka‘i has almost as many with 494 species, while Lāna‘i has 362 species. The smaller and relatively arid islands of Ni‘ihau and Kaho‘olawe have only 99 and 68 native species, respectively. The Northwestern Hawaiian Islands together are home to 52 native species. If the four islands comprising Maui Nui are considered together, there are 705 species. Looking at the number of single-island endemics, defined as those endemic Hawaiian species whose natural range is limited to a single island, Kaua‘i is the clear winner with 219 species, far more than any other island. O‘ahu is second in single-island endemics with 140 species, followed by Maui with 89 species and Hawai‘i with 87 species. The relatively low number of single-island endemic species on Hawai‘i, despite its large size and topographic heterogeneity, is likely due to its youthful age, which means that there has been insufficient time for evolution to fully saturate available niches with selected genotypes. The low level of single-island endemics on Maui is related instead to its former land connection with Lāna‘i and Moloka‘i as Maui Nui. The number of single-island endemics is 36 on Moloka‘i and 8 on Lāna‘i, while Ni‘ihau and Kaho‘olawe have 2 and 3 species, respectively. Maui Nui considered as one island has 204 single-island species present, still lower than Kaua‘i.

ur knowledge of the size and diversity of the Hawaiian flora remains in a state of modest dynamic change as new species continue to be uncovered and species concepts modified. The continual discovery of new species is remarkable for such a well-studied and carefully collected flora. Many of the newly described species in recent years have been found in virtually inaccessible areas of vertical cliffs, particularly on Kaua‘i, where plant collectors have risked life and limb by hanging from climbing ropes. In talking about the Hawaiian flora, we need to define several terms that characterize groups of species before we talk about species richness. Native species are those species that are present in the flora due to natural processes of dispersal and evolution. We further divide the native species into two groups. Indigenous species are those that occur naturally in the Hawaiian Islands but also have a biogeographic range of natural occurrence outside of the Islands. Endemic species are those that occur only in the Hawaiian Islands and nowhere else, which indicates that they have likely evolved in situ into new species from their ancestral colonizer. In contrast, alien or nonnative species were brought to the Hawaiian Islands either intentionally or unintentionally by human activities. Some of these came with the Polynesians, such as coconuts and yams (chapter 6), while others were introduced more recently by colonists from North America, Europe, and Asia. This latter group includes some of the most harmful alien species in the Islands (chapter 7). The native vascular plant flora of the Hawaiian Islands at the time of this writing includes 1,207 species, comprising 163 ferns and fern allies, 140 monocots, and 904 dicots. A remarkable feature of this flora is a level of endemism that is unparalleled anywhere else in the world. For the total native flora, 88% of species are restricted in distribution to the Hawaiian Islands, with rates of endemism particularly high in the dicots at 93%. Monocots and ferns with fern allies each have lower levels of endemism at 73% and 75%, respectively. A number of endemic subspecific taxa (i.e., varieties and subspecies) have also been identified for the Hawaiian flora where the overall species range of distribution occurs beyond the Islands. If these taxonomic levels are included in floristic totals, the native vascular plant flora rises to a total of 1,352 named taxa. Looking at data on Hawaiian floristics at the level of individual islands, Maui and Kaua‘i are tied in having the greatest number of species with 629. O‘ahu is next in diversity with 587 species (table 2.1). Hawai‘i is significantly lower in richness

Colonization and Adaptive Radiation The Hawaiian Islands form one of the most isolated land masses in the world. It is not surprising, therefore, that successful plant colonization of the Islands has been a rare event. There are thought to be about 375 colonizing lineages for vascular plants that have evolved into the current native flora. If we use the age of Kaua‘i at about 4.7 Ma as a starting point, that means that there has been a net average of about one successful colonization event every 12,500 years. However, we know from molecular studies, as described later, that there are a few Hawaiian lineages with an evolutionary history that predates the existence of any of the modern islands. These are lineages that hopped from island to island down the Northwestern Hawaiian chain, dating back perhaps as long ago as the geological formation of Kure at the northwestern end of the chain. It is clear, however, from geological studies that only low island masses existed between the submergence of Kōkō Seamount about 33 Ma and the formation of Lisianski Island about 23 Ma. The movement of terrestrial organisms along the

19

20

PLANT DIVERSITY AND EVOLUTION

chain during this period must have been restricted to lowland species. The genus Hillebrandia (Begoniaceae) and the lobeliads (Campanulaceae) are among the few plant lineages that appear to predate the origin of Kaua‘i. Fruitflies (Drosophila) at 26 Ma and damselflies at 9.6 Ma are other lineages that evolved as a lineage before the existence of the modern islands. How were these ancestral colonists able to get to the Hawaiian Islands? We can get quite a good idea of this by looking at the morphological and physiological traits of seeds and fruits of the modern Hawaiian angiosperm flora. For indigenous species, the best guesses are that about one-third of the species arrived by rafting or in ocean drift. This correlates well with the coastal habitats in which many of these lineages are found today. The other two-thirds of the indigenous species arrived through the action of birds, either as seeds stuck to feathers or feet or as ingested seeds or fruits. For lineages that produced endemic species, the role of birds is thought to be much more important, accounting for about 83% of the successful colonizations, with only 17% due to ocean dispersal. For ferns and fern allies, most species likely came from atmospheric transport of their tiny spores. Several molecular and morphological techniques have been utilized by plant systematists to provide a fairly accurate assessment of the number of original colonizing species that gave rise to the modern Hawaiian flora. When these colonizing lineages are divided into groups based on the level of resulting speciation, it becomes clear that widespread adaptive radiation in the Hawaiian Islands has not been common. Nearly two-thirds of these successful colonizing lineages occur today with only a single Hawaiian species. Just over half of these single-species lineages are widespread indigenous plants that did not undergo speciation after reaching the Hawaiian Islands. The remaining colonizers evolved to become a single endemic species, without any further speciation.

Flowering Plants The current total of 1,044 native angiosperm species in the Hawaiian Islands adds 88 species to those reported in The Manual of Flowering Plants of Hawai‘i in 1990, with most of the additions based on newly described taxa. Of these new species, 82 are dicots and 6 are monocots. At the species level, 90% of the native flowering plant species are endemic to the Hawaiian Islands. This is a remarkably high level of endemism for any flora. Endemism is also quite high compared to other floras at the generic level, although new studies have been slowly reducing the number of genera considered to be endemic to the Hawaiian Islands, as molecular data show close affinities with groups more broadly distributed. The diversity of endemic species in the Hawaiian flora is not equally distributed across genera and families. Much of the endemic flora has resulted from the extensive speciation that has occurred in a small number of highly successful lineages. The 5 largest families together form 40% of the Hawaiian angiosperm flora (table 2.2). This group is led by the Campanulaceae (141 species), followed by the Asteraceae (98 species), Lamiaceae (61 species), Rubiaceae (61 species), and Gesneriaceae (59 species). There are 13 families with more than 20 species. Only the Cyperaceae among the 13 largest families has an endemism level of less than 50%.

Even more remarkable is how few colonizing lineages evolved to produce the species in these 5 largest families. Just 29 founding lineages are thought to have become established, and they radiated into the 411 species present today in these families. Most successful of all of these founding lineages, as described below, are the lobeliads that make up the 141 native species of Campanulaceae. These are all thought to be evolved from a single colonizing lineage. The next largest lineages arising from single colonizers are the endemic genera of Hawaiian mints (Lamiaceae), Cyrtandra (Gesneriaceae), Melicope and Platydesma (Rubiaceae), and Schiedea (Caryophyllaceae). There are 5 Hawaiian genera with 30 or more species. Leading this group is Cyanea (Campanulaceae) with 79 species, followed by Cyrtandra (Gesneriaceae) with 59 species, Melicope (Rutaceae) with 48 species, Schiedea (Caryophyllaceae) with 34 species, and Phyllostegia (Lamiaceae) with 32 species. Because of the speciation that has occurred in the major lineages in these large genera, endemism is 100% in these 5 groups. There are 5 additional genera with more than 20 species: Dubautia (26 species), Peperomia (25 species), Clermontia (22 species), Stenogyne (22 species), Kadua (22 species), and Pritchardia (22 species). All of these are endemic with the exception of 2 species of Peperomia.

LOBELIAD ALLIANCE A casebook example of adaptive radiation in an Island lineage is the lobeliad alliance, consisting of the genera Clermontia, Cyanaea, Delissea, Brighamia, Lobelia, and Trematolobelia, with a total of 141 Hawaiian species. Although once thought to have had their origin derived from 3 to 5 independent colonization events, molecular data now indicate that the group has arisen from a single colonizing species. Thus, this single lineage has produced 12% of the entire native Hawaiian flora. The ancestral colonizer is estimated to have arrived about 13 Ma as a woody, wind-dispersed, bird-pollinated species adapted to open habitats at middle elevations. This age of colonization is far older than the age of Kaua‘i and thus must have occurred on one of the Northwestern Hawaiian Islands. The morphological and ecological divergence of the Hawaiian lobeliads from their ancestral founder is remarkable. Growth forms within this lineage include trees, treelets, shrubs, stem succulents, and bog rosette plants. Leaf shape, floral morphology, and modes of pollination and seed dispersal similarly vary across species. Flower form has coevolved with native Hawaiian honeycreepers. Native habitats for lobeliads include dry forests, mesic forests, rainforests, open bogs, and sea cliffs, with light environments covering the entire range from full sun to shaded forest understory. The invasion of closed canopy wet forests by Cyanea, the largest genus with 79 species, appears to have been associated with accelerated rates of speciation. Most taxa in this genus are single-island endemics, with each of the higher islands having a similar number of species. This pattern suggests that some genera may diversify and speciate relatively rapidly to fill available niches on a single island.

HAWAIIAN MINTS Another group with an impressive level of speciation in the Hawaiian Islands is formed by 3 genera of endemic Hawaiian mints. These genera, with 59 species, have evolved from a

a

b

d

c

e

2.1  Diversity

of growth form in Hawaiian lobeliads: a) Brighamia insignis; b) Lobelia gloria-montis; c) Trematolobelia macrostachys; d) Cyanea hamatiflora; e) Clermontia

arborescens

2.2 

‘I‘iwi on Lobelia grayana

2.3 

Fleshy fruit of Phyllostegia grandiflora

22

PLANT DIVERSITY AND EVOLUTION

single colonizer from western North America about 5 Ma. Haplo­ stachys, with 5 known species, 4 of which are now extinct, once occurred more widely in relatively dry habitats at low to middle elevations. The one surviving species is restricted to the saddle area between Mauna Kea and Mauna Loa on Hawai‘i. The fruits of Haplostachys are dry nutlets, an ancestral trait in comparison to the fleshy fruits of the other two genera. Phyllostegia includes 32 endemic Hawaiian species, most commonly growing in mesic to wet forest habitats on all of the major islands. One species, now presumed extinct, once occurred on Kure and Midway atolls and Laysan Island. Two additional species have been described, 1 from Tahiti and 1 from Tonga. The third genus, Stenogyne, is a relatively recent evolutionary lineage that includes 22 endemic species. These taxa are found on all of the higher islands but with greater speciation on Maui and Hawai‘i. Floral structure in Sten­ ogyne has become adapted for pollination by endemic Hawaiian honeycreepers, a change from insect pollination in Haplostachys and Phyllostegia. This pollination syndrome, once produced, restricted possibilities for gene flow with these 2 genera.

HAWAIIAN ASTERACEAE The Asteraceae represent a diverse and well-studied group in the Hawaiian flora, with all but 1 of the 98 native species endemic to the islands. Four of the estimated 10 founding ancestors of this family have given rise to lineages with major speciation. This has occurred in the silversword alliance, the sixth largest Hawaiian lineage, and in Bidens, Melanthera, and Tetramolopium. In each of these lineages there has been broad morphological and ecological differentiation despite a relatively low level of genetic divergence. The silversword alliance, with 3 endemic genera—Argyroxi­ phium, Dubautia, and Wilkesia—presents a particularly impressive story of adaptive radiation, with 33 endemic species evolved from a single ancestral colonizing lineage closely related to the California tarweed Carlquistia muirii. Dubautia forms the largest of these genera, with 26 species. Molecular clocks suggest that the silversword alliance first became established on a higher Kaua‘i or older island about 5.2 Ma. Interestingly, these molecular data show Argyroxiphium as the ancestral group in the alliance despite its restriction today to Maui and Hawai‘i. Adaptive shifts in growth form and ecophysiological traits as well as ancient hybridization between species have been major factors in

a

2.4 

b

c

the diversification of this group. From an herbaceous perennial ancestor, there has been the evolution of shrub, rosette, liana, and tree growth forms, with both polycarpic and monocarpic reproductive systems present. Shifts from wet to dry habitats have occurred in each of the primary lineages of the alliance, and on each of 3 and possibly all 4 of the major island masses of Kaua‘i, O‘ahu, Maui Nui, and Hawai‘i. Nineteen endemic species have evolved in Bidens from a single colonizer thought to have arrived from Latin America. Morphological diversity in the Hawaiian Bidens is far greater than that present in the nearly 200 other species of the genus. Not only are there both herbaceous and woody taxa, but also there is major adaptive variation in reproductive morphologies, with diversity in sexual expression of flowers to promote outcrossing. This variation has included the widespread development of dioecy, shifts in at least one case from wind to bird pollination, and a shift from biotic to abiotic dispersal of seeds. Much less is known about evolution of the 15 endemic species of Hawaiian Melanthera, once considered to be a subgenus of Lipochaeta. Tetramolopium presents an interesting example of a relatively recent colonization and adaptive radiation in the Hawaiian Islands. The center of diversity of the genus is in subalpine and alpine habitats of New Guinea. Outside of New Guinea, there are 11 species in the Hawaiian Islands and 1 in the Cook Islands, which is a range extension of 1 of the native Hawaiian species. The basal taxon of Tetramolopium in the Hawaiian Islands is T. humile, a subalpine and alpine subshrub known from Maui and Hawai‘i. This species and most of the New Guinea species of Tetramolopium have glandular trichomes on their fruits, making them relatively easily dispersed on the feathers of birds. Molecular data suggest that Tetramolopium first colonized Maui in the Pleistocene, no more than 1.5 Ma, becoming established in subalpine and alpine regions much like the habitat of its New Guinea progenitors. Despite evidence that speciation began no more than 600,000–700,000 years ago, Tetramolopium has undergone remarkable morphological and physiological radiation to allow its derived species to adapt to habitats from sea level to 3,000 m elevation and develop growth forms such as low alpine subshrubs, prostrate coastal shrubs with succulent leaves, and forest shrubs up to 2 m tall. Tetramolopium is notably absent, however, from rain forest habitats, suggesting the possibility of competitive

d

Silversword complex: a) Dubautia ciliolata; b) Wilkesia gymnoxiphium; c) Carlquistia muirii (California); d) Argyroxiphium sandwicense subsp. macrocephalum

PLANT DIVERSITY AND EVOLUTION

a

b

c

d

2.5 

Diversity of leaf form in Metrosideros polymorpha: a) var. glaberrima; b) var. incana; c) var. pumila; d) var. pseudorugosa

exclusion by older lineages. The presence of T. rockii on coastal deposits no more than 15,000 years in age on O‘ahu and Moloka‘i suggests very recent evolution of this taxon. The earliest colonization of Tetramolopium on Maui and evidence of multiple back colonizations to Moloka‘i and O‘ahu sharply contrast with the common patterns of west-to-east migration patterns present in the evolution of most Hawaiian plant and animal groups. While molecular studies have generally supported conclusions from previous evolutionary hypotheses based on morphological data, there are sometimes surprises. One such unexpected result has come from molecular studies of the endemic Hawaiian genus Hesperomannia, another member of the Asteraceae. Previously this small genus of 3 tree species was thought to have its closest relationships with several genera of the tribe Mutiseae in South America. DNA phylogenies now show that Hesperomannia is closely allied to African members of the tribe Vernonieae.

METROSIDEROS (MYRTACEAE) The excellent dispersal ability of the tiny wind-dispersed seeds of Metrosideros in the Myrtaceae family has led the group to be described as a super-tramp lineage. The genus presents an interesting example of adaptive plasticity in morphology and ecophysiological traits with little speciation. This trait of dispersibility has allowed a lineage of Metrosideros from New Zealand to colonize high volcanic islands across the Pacific Basin, while being absent from atolls and raised limestone islands. Metrosideros is thought to have colonized the Hawaiian Islands in the Middle to Late Pleistocene, about 1.1 Ma, from a

staging point in the Marquesas Islands. Traditional taxonomy has divided Metrosideros into 5 species in the Hawaiian Islands, with M. polymorpha being the most widespread and dominant over mesic and wet forests on all of the larger islands. As its specific name suggests, M. polymorpha exhibits a remarkable degree of ecological and morphological plasticity. It can be found from sea level to treeline and from dry forests to the wettest rain forests. Eight varieties of M. polymorpha have been named, with leaf pubescence on the lower surface and leaf form and size among the primary distinguishing characteristics between these varieties. Despite a number of recent molecular studies, it remains unclear whether the morphological variation within Hawaiian Metrosideros is due to hybridization, genetic polymorphism, phenotypic plasticity, or some combination of these processes. Ongoing molecular studies are helping to better resolve the relationships of the Hawaiian taxa and may well result in a revision of species concepts. Studies of Metrosideros across an elevational gradient on Mauna Loa have helped explain the selective significance of these differences in leaf traits. In this gradient, there are 5 varieties of M. polymorpha. The varieties incana and polymorpha have pubescent leaves, with the former at low to middle elevations and the latter at middle to high elevations. Two other varieties, macro­ phylla and glaberrima, have glabrous leaves and occur at middle and middle to high elevations, respectively. Pubescent varieties that dominate the early stages of tree succession at arid lower elevations and on recent lava flows have higher growth rates, greater tolerance of water stress, and higher fecundity than glabrous

23

24

PLANT DIVERSITY AND EVOLUTION

varieties. There appears to be high genetic polymorphism for leaf traits among populations, with physiological properties linked to these foliar morphologies and influencing the observed patterns of elevational and successional patterning of populations and perhaps incipient speciation. The 4 other varieties of M. polymorpha are restricted geographically and ecologically to specialized habitats. The var. dieteri is a small tree with glabrous large leaves known only from wet forests on Kaua‘i, and var. newellii is a glabrous shrubby tree restricted to stream sides on the island of Hawai‘i. The final 2 varieties are shrubby bog specialists. These are var. pumila on Kaua‘i, Moloka‘i, and Maui, and the small-leaved var. pseudoru­ gosa known only from West Maui. Both of these bog varieties generally have pubescent leaves, although this trait is variable in var. pumila.

OTHER LINEAGES Cyrtandra (Gesneriaceae) is a large genus of up to 600 species, extending in range from Southeast Asia and New Guinea across the Pacific Basin, with 59 endemic Hawaiian species. There are convincing molecular data suggesting that all of these Hawaiian species have evolved from a single colonizing ancestor, making this genus tied with the Hawaiian mints as the second largest endemic lineage. Despite this extensive speciation, there has been relatively little ecological differentiation, with most species growing as understory shrubs or small trees with white flowers and fleshy berries and being restricted in habitat to wet forest habitats. Across the Pacific Islands, most species of Cyrtandra are narrowly distributed endemics occupying only a single archi­ pelago, a single island, or even a single valley. A large number of species of Melicope (Rutaceae) in the Hawaiian Islands have evolved from a single colonizing ancestor. Recent molecular studies have indicated that the 4 Hawaiian species of Platydesma, typically considered an endemic genus, are nested within Melicope, and both groups are derived from a single colonizing ancestor, making this the fourth largest lineage for the islands with 52 species. Melicope is a genus of about 150 species extending across the Australasian region, with the Hawaiian Islands a notable center of speciation. The 34 Hawaiian species of Schiedea (Caryophyllaceae) form the fifth largest lineage from a single ancestor. Most species are single-island endemics. If Moloka‘i, Lāna‘i, and Maui are considered as part of the single Maui Nui landmass, only 4 species occur on more than a single island. Thus, Schiedea provides a significant model system for understanding speciation resulting from isolation on different islands compared to allopatric speciation, which results from ecological habitat shifts within a single island. Interisland colonization and speciation has moved from older to younger islands, with sufficient isolation occurring to form single-island endemics. Closely related species pairs occurring on the same island tend to differ in habitat, and thus they appear to be ecologically isolated. Speciation in Schiedea is associated with variations in breeding system, pollination system, and habitat. Dioecy has evolved multiple times within the genus, and there is a wide diversity of breeding systems. Two sections of species in Schiedea have evolved floral nectaries, a morphology unique among the Caryophyllaceae, apparently as a coevolved trait with Hawaiian bird pollinators. Also unusual among related groups of Caryophyllaceae is the woody growth habit of many

species, a characteristic example of secondary evolved woodiness in island floras. Some of the Hawaiian evolutionary lineages are clearly ancient and far exceed the age of the major islands. An extreme example of this can be seen with Hillebrandia sandwicensis, a monotypic genus in the Begoniaceae known only from wet forests on Kaua‘i, Moloka‘i, Maui, and with a record of an early observation on O‘ahu. Molecular data suggest that this genus is close to basal within its family and likely evolved 51–65 Ma. Thus, it has survived as a relict by island-hopping down the Hawaiian chain to younger islands as they have formed. Not all speciose genera in the Hawaiian flora have evolved from a single colonizer. A notable example of a more complex origin lies with the genus Scaevola (Goodeniaceae), with 10 species, 9 of which are endemic. Molecular data have verified earlier morphological studies establishing that these species are the result of 3 independent colonizers. One of these is the indigenous S. taccada, a widespread species on coastal shores across the Pacific and Indian oceans. The endemic tetraploid S. glabra represents a long-distance dispersal of this lineage from Australia. The other 8 endemic diploid species are part of a clade that includes wide-ranging taxa. It is notable that 3 of the 6 successful events of establishment of Scaevola taxa away from Australia are in the Hawaiian flora, despite the fact that no new species have evolved on continental areas outside of Australia. Even in small genera, there are examples of multiple colonization events. This can be seen in Santalum (Santalaceae), with 6 endemic species evolving from 2 independent colonizations. The first dispersal event came from Australia about 1.0–1.5 Ma, while the second, also from Australia, occurred about 400,000– 600,000 years ago.

Ferns and Fern Relatives The Hawaiian flora contains 163 species of native ferns and fern allies, with 122 of these species endemic (75%) and 41 indigenous. This adds 2 species to the total number of native species given in Palmer’s Hawai‘i’s Ferns and Fern Allies (University of Hawai‘i Press, 2003), although some of the generic concepts have changed since this book. Two fern genera are recognized as endemic to the Hawaiian Islands. These are Adenophorus (Polypodiaceae, formerly separated into the Grammitidaceae) and Sadleria (Blechnaceae), each of which has evolved into multiple species from a single colonizing ancestor. Formerly, Diellia was recognized as an endemic genus, but these species are now placed within Asplenium. Despite a relatively high diversity of ferns and fern allies at the generic level, 5 genera account for 41% of the native species. These are Asplenium (27 species), Dryopteris (11 species), Huper­ zia (10 species), Adenophorus (10 species), and Elaphoglossum (9 species). In contrast to the pattern seen in the flowering plants, where large species groups have evolved from a single colonizing lineage, only Adenophorus of these large genera represents speciation from a single lineage. The current estimate is that the 27 native species of Asplenium have developed from 18 separate colonizing lineages. Similarly, the 9 native species of Elaphoglossum likely represent 6 colonizing lineages, while the native species each of Dryopteris and Huperzia appear to have developed from 5 colonizing lineages each.

PLANT DIVERSITY AND EVOLUTION

a

b

across the equator in association with seasonal shifts in the Intertropical Convergence Zone (ITCZ). While we do not think of ferns as plants that readily invade and become established in areas outside of their natural range, the Hawaiian Islands has provided a home for at least 33 naturalized fern species. Two of these have become major invaders of native forest habitat. Sphaeropteris cooperi (Australian tree fern, Cyatheaceae) has become a serious invader of pristine wet forest areas on the windward slopes of Kaua‘i (see chapter 7). Angiop­ teris evecta (Marattiaceae) has also become an aggressive alien invader, becoming well established in wet forests in shaded valleys and mountain ridges on Kaua‘i, O‘ahu, Lāna‘i, and Maui.

Lower Plants and Fungi 2.6  Endemic genera of Hawaiian ferns: a) Adenophorus epigaeus (Polypodiaceae); b) Sadleria unisora (Blechnaceae)

It seems logical that colonizing lineages among the ferns and fern allies have not developed adaptive radiations to the degree seen in flowering plants. The ease of movement of small wind-dispersed fern spores limits possibilities for genetic divergence between populations—and thus speciation. Approximately 60% of Hawaiian ferns and fern allies are present on all 6 of the largest islands, excluding Ni‘ihau and Kaho‘olawe, demonstrating the relatively strong dispersal ability of fern spores. Not surprising also is the fact that the number of species of native fern and fern allies present on the major islands is remarkably similar. Kaua‘i has 136 native species, with 13 endemic species restricted to this island, and O‘ahu has 130 species, with 5 single-island endemics. Moloka‘i has 121 native species, while Lāna‘i has 108 native species. Neither of these islands has a single-island endemic species. Maui has the largest number of native fern species with 138, with 2 single-island endemic species. Hawai‘i has 131 native fern species, but none of these is restricted to the island. Ferns are relatively rare and low in diversity on the dry lower islands. Ni‘ihau has just 6 reported fern species, and Kaho‘olawe has 5 species. If Maui Nui is considered as a single island, then it leads in fern diversity with 141 species. Hawaiian ferns have received less phylogenetic and biogeographic study than have Hawaiian angiosperms. Nevertheless, there are now molecular phylogenies of several Hawaiian fern genera that offer some insight into the biogeographic patterns of dispersal and speciation in these groups. Unlike flowering plants, where propagules have been dispersed to the Islands largely by birds and flotation, the tiny spores of ferns and fern allies are readily dispersed in wind currents. Spore dispersal from the Indo-Pacific region by the northern subtropical jet stream is strongly supported for 2 of the Hawaiian lineages of Polystichum, 3 of the 5 endemic lineages of Dryopteris, and possibly the endemic Grammitis hookeri clade. The endemic genus Adenopho­ rus has neotropical relationships and thus likely reached the Hawaiian Islands from the trade winds or perhaps chance dispersal in storms. Stenogrammitis (Polypodiaceae) is another lineage in the Hawaiian Islands with neotropical ancestry, suggesting dispersal by either the trade winds or possibly from the South Pacific

There is still much to learn about the bryophytes, lichens, and fleshy fungi of the Hawaiian Islands, but we know enough to have some idea of their diversity. Bryophytes can be divided into 3 taxonomic groups: the mosses, hornworts, and liverworts. There are 241 native species of mosses reported for the Hawaiian Islands, with 75 of these endemic (31%). Just 1 of the 4 native species of hornworts is endemic. Diversity and endemism is relatively high in the liverworts, with 174 native species and 102 of these endemic (59%). Lichens represent symbiotic relationships between fungi and algae, with the scientific name referring to the fungal component. Most of these fungal components are ascomycetes. There are 880 taxa of lichens presently reported for the Hawaiian Islands. Macrofungi—large, easily seen species of basiomycetes and nonlichenized ascomycetes—remain relatively poorly studied in the Hawaiian Islands. Considering fleshy fungi with the basiomycete order Agaricales, recent studies have listed 310 species distributed over 83 genera and 14 families. Many of these are nonnative to the Hawaiian Islands, having been brought in with introduced plants, but about half of these species are considered endemic.

Traits of Adaptive Radiation The endemic flora of the Hawaiian Islands, as in many other archipelagos, exhibits a number of distinctive adaptive traits in comparison to their colonizing continental ancestors. These are traits that are hypothesized to facilitate the successful colonization of open and unsaturated ecological habitats. The term adap­ tive radiation is usually defined as the diversification of a lineage into species that exploit a variety of different resources or habitats and that differ in the morphological and ecophysiological traits that allow them to exploit these new resources or habitats. Often lineages with no evolutionary relationship evolve similar traits as a result of adapting to similar environments or ecological niches. This process is called convergent evolution.

HYBRIDIZATION Hybridization has often been suggested as a key element of speciation on island archipelagos, although it is not clear that hybridization is more frequent on oceanic islands than in continental regions. Hypothetically, the formation of hybrids can lead to the establishment of new stabilized taxa with novel or intermediate morphological or ecophysiological traits. The success of

25

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PLANT DIVERSITY AND EVOLUTION

a

b

c

d

Hybridization in Hawaiian silversword: a) Argyroxiphium sandwicense subsp. macrocephalum; b) Dubautia menziesii; c) hybrid with stalked rosette; d) hybrid with branched inflorescence and yellow flowers

2.7 

new hybrids would be expected to be increased in the presence of open or unsaturated habitats, a condition present in oceanic archipelagos with a dynamic geologic history such as the Hawaiian Islands. Natural hybrids are widely present in the Hawaiian flora in many genera. Because genetic diversification often lags behind morphological differentiation in the Hawaiian flora, interspecific hybridization is common. Cyrtandra forms the most prominent example, with at least 67 interspecific hybrids identified. Hybridization is also well described in Dubautia, where hybrid populations and introgression have been studied in detail between Dubautia arborea and D. ciliolata and between D. scabra and D. ciliolata on Hawai‘i. Other examples of interspecific hybridization can be seen in Bidens, Euphorbia (Chamaesyce), Clermontia, Coprosma, Cyanea, Melanthera, Psychotria, Schie­ dea, Stenogyne, Tetramolopium, and Wikstroemia. Hybridization is also present in Hawaiian ferns, as with natural hybrids between Adenophorus hymenophylloides and both A. tamariscinus and A. tripinnatifidus. Intergeneric hybridization also occasionally occurs in Hawaiian lineages, as seen in hybrids between Argyroxiphium sandwicense subsp. macrocephalum and Dubautia menziesii on

Haleakalā, Maui. It has been hypothesized that the endemic genus Lipochaeta originated from an intergeneric hybridization between Melanthera and an unknown taxon, perhaps Wedelia. A more unusual case of intergeneric hybridization can be found in rare hybrids discovered on Kaua‘i between the native fern Lind­ saea ensifolia and the introduced Sphenomeris chinensis. The oddly shaped fronds of the hybrids are intermediate in morphology between the quite different fronds of the 2 parents.

SEXUAL DIMORPHISM Successful establishment on remote oceanic islands is clearly a rare event, and thus new island plants often suffer from low levels of genetic diversity. It is logical then that species with mechanisms to promote outcrossing may be more able to adapt to varied conditions and speciate. The selective forces favoring sexual dimorphism are those favored by the promotion of outcrossing mechanisms. More than 20% of the native flowering plants in the Hawaiian Islands have sexually dimorphic flowers, a level much higher than present in continental areas. These dimorphic flowers include species with complete dioecy, forming separate male and female plants, as well as intermediate forms (gynodioecy and polygamodioecy) with a mix of perfect and single-sex flowers.

PLANT DIVERSITY AND EVOLUTION

a

b

2.8 

Sexual dimorphism in Coprosma ernodeoides: a) male; b) female

There is a second reason why outcrossing may be favored in island groups. Dioecious species commonly exhibit altered patterns of resource allocation between male and female plants, with female plants requiring greater resource investment in reproduction than male plants because of the costs of fruit production. Lower investment costs may allow male plants to colonize resource-poor environments where female plants would not be successful, with female plants localized in areas of higher resource availability. The presence of dioecy in Hawaiian plant lineages has been shown to be positively associated with the presence of fleshy fruits, woody growth form, and small, inconspicuous flowers. Selection for sexual dimorphism occurs not only in many successful colonizing plants, but has also evolved in situ. At least 12 colonizing lineages in the Hawaiian Islands developed sexual dimorphisms after arrival, and these lineages gave rise to about one-third of the sexually dimorphic species in the flora. An interesting example of this can be seen in Schiedea, where there have been multiple independent origins of sexual dimorphism.

INSULAR WOODINESS One of the interesting traits of island floras is that many flowering plant families and genera that typically are comprised of herbaceous species have secondarily evolved woody growth forms as shrubs or trees. This phenomenon was noted back in the middle of the nineteenth century by Charles Darwin and Alfred Russel Wallace in their field studies. Good examples of the evolution of insular woodiness in colonizing lineages can be seen in

2.9 

Insular woodiness in Euphorbia olowaluana

Hawaiian plant families such as the Amaranthaceae (Chenopo­ dium), Asteraceae (Dubautia, Hesperomannia), Campanulaceae (Clermontia, Cyanea), Schiedea (Caryophyllaceae), Euphorbiaceae (Euphorbia section Chamaesyce), Plantaginaceae (Plantago), and Violaceae (Viola). This insular woodiness is generally suggested to have evolved as a result of a combination of release from competitive interactions and absence of large herbivores in the diverse and evolutionarily unsaturated environments of isolated island ecosystems. Another hypothesis is that this trait may relate to selective pressures promoting outbreeding in island floras. Pollination pressures may promote selection for larger plants with greater longevity and conspicuous inflorescences to increase the likelihood of successful outcrossing. Under this hypothesis, the specific woody growth form evolved is unrelated to climate conditions. Such a mechanism has been hypothesized to explain the diversity of woody growth forms that have evolved in species of Echium (Boraginaceae) in the Canary Islands from an herbaceous colonizing lineage. Some of these assumptions on the role of convergent evolution in the development of large woody island lineages have been called into question by molecular studies of the giant lobeliads of the Hawaiian Islands and East Africa, together with other large lobeliad species from eastern Brazil,

27

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PLANT DIVERSITY AND EVOLUTION

French Polynesia, and Southeast Asia. These lobeliads all appear to be derived from an ancestral lineage that was woody and probably African.

LOSS OF HERBIVORE DEFENSES Herbivores often represent a strong selective factor on the growth and survival of plants, and thus it is not surprising that selective responses to herbivory have resulted in the widespread evolution of chemical plant toxins, aromatic oils, and morphological features such as spines, prickles, and dense hairs to reduce pressures of herbivory. However, there are a number of Hawaiian plant lineages that appear to have lost such ancestral traits, hypothetically because of an environment that lacks terrestrial mammals that are typically the primary herbivores in most areas of the world. An example of loss of chemical defenses is found in the Hawaiian mints (Lamiaceae), where none of the 58 endemic species retains aromatic oils in its foliage, which are thought to be deterrents to herbivores. While this seems a reasonable explanation, this is not a hypothesis that can be easily tested. A loss of protective chemicals is certainly not universal in the Hawaiian flora. Wikstroemia, which also grows outside of the Hawaiian Islands, is notably poisonous, and Hawaiian species retain sufficient toxin to have been used by the Polynesians to poison fish in tide pools. Although not formally tested, the milky saps of endemic species of Euphorbia are likely toxic as well. Many authors have pointed to the relative absence of thorns and spines in the Hawaiian flora as an example of the reduced pressure from vertebrate herbivores. For example, the 2 Hawaiian species of Rubus have their prickles reduced to soft deciduous structures with no defensive function. Loss of protective function also seems evident in many Hawaiian species of Urticaceae, where several endemic species lack the stiff hollow hairs that normally characterize the family. Two examples of endemic Hawaiian species that retain ancestral traits of sharp prickles are Erythrina sandwicensis and Argemone glauca, perhaps reflecting more recent arrival in the Hawaiian Islands.

2.10 

Absence of sharp prickles in Rubus hawaiiensis

There is another Hawaiian plant group where prickles appear to have evolved in situ, rather than being an ancestral trait. These are the Hawaiian lobeliads, where about 20 species of Cyanea representing 4 distinct lineages from the founding population have evolved prickles. These prickles, some of which may be sturdy structures up to 10 mm in length, are most abundant

and best developed on the lower stems and young leaves of the plants. It has been suggested that these prickles evolved to reduce herbivory by now extinct flightless birds that once occurred in the Hawaiian Islands. The birds that might well have played this role were 4 species of flightless geeselike ducks, known as moa nalo, that once occurred widely.

2.11 

Prickles on the stem of Cyanea horrida

It has also been hypothesized that heterophylly in species of Cyanea—that is, species where the morphology of juvenile leaves is quite different from that of adult leaves—also represents an adaptation to lower herbivory by visually orienting flightless birds. Interestingly, the traits of juvenile prickles and heterophylly are often, but not invariably, correlated in Cyanea and increase in frequency with younger island ages. Prickles are absent in all of the 16 species of Cyanea on Kaua‘i, and they increase steadily from 5 of 14 species on O‘ahu to 11 of 26 species on Maui Nui and finally to 6 of 12 species on Hawai‘i.

2.12 

Tubercles on young plants of Cyanea tritomantha

PLANT DIVERSITY AND EVOLUTION

TABLE 2.1.  Distribution of native species, endemism, and evolutionary lineages for the native flora on the individual Hawaiian Islands (DATA TAKEN FROM APPENDIX: NATIVE VASCULAR PLANTS OF THE HAWAIIAN ISLANDS)

TOTAL VASCULAR PLANT FLORA Species

Endemism (%)

Single-island endemic

Lineages present

Species/lineage ratio

Northwestern Hawaiian Islands

52

50

5

48

1.08

Ni‘ihau

99

49

2

88

1.13

Kaua‘i

629

81

219

324

1.94

O‘ahu

587

79

140

322

1.82

Moloka‘i

494

76

36

305

1.62

Lāna‘i

362

75

8

249

1.45

Maui

629

80

89

338

1.86

68

57

3

58

1.17

Maui Nui

705

82

204

293

2.41

Hawai‘i

520

77

87

319

1.63

TOTAL

1,207

88

589

376

3.21

Species

Endemism (%)

Single-island endemic

Lineages present

Species/lineage ratio

Northwestern Hawaiian Islands

37

54

5

34

1.09

Ni‘ihau

70

53

2

63

1.11

Kaua‘i

405

87

188

168

2.41

O‘ahu

381

86

130

172

2.22

Moloka‘i

298

82

33

158

1.89

Lāna‘i

205

80

7

130

1.58

Maui

399

86

82

171

2.33

50

58

3

45

1.11

Maui Nui

463

88

186

176

2.63

Hawai‘i

307

84

79

155

1.98

Total

904

93

529

192

4.70

Kaho‘olawe

ANGIOSPERMS: DICOTS

Kaho‘olawe

29

30

PLANT DIVERSITY AND EVOLUTION

ANGIOSPERMS: MONOCOTS Species

Endemism (%)

Single-island endemic

Lineages present

Species/lineage ratio

Northwestern Hawaiian Islands

14

43

0

13

1.08

Ni‘ihau

23

39

0

10

1.21

Kaua‘i

88

67

18

64

1.38

O‘ahu

76

59

5

59

1.27

Moloka‘i

75

64

3

58

1.29

Lāna‘i

49

69

1

36

1.36

Maui

92

65

4

68

1.35

Kaho‘olawe

13

62

0

8

1.63

101

66

16

71

1.42

82

61

8

65

1.26

140

73

39

79

1.78

Endemism (%)

Single-island endemic

Lineages present

Species/lineage ratio

Maui Nui Hawai‘i Total

FERNS AND FERN ALLIES Species Northwestern Hawaiian Islands

1

0

0

1

1.00

Ni‘ihau

6

50

0

6

1.00

Kaua‘i

136

71

13

92

1.48

O‘ahu

130

70

5

91

1.43

Moloka‘i

121

69

0

89

1.36

Lāna‘i

108

69

0

83

1.30

Maui

138

72

3

99

1.39

5

40

0

5

1.00

Maui Nui

141

72

2

101

1.40

Hawai‘i

131

69

0

99

1.32

TOTAL

163

75

21

104

1.56

Kaho‘olawe

PLANT DIVERSITY AND EVOLUTION

TABLE 2.2.  Species richness and endemism in Hawaiian angiosperm families with more than 20 species

Species in Hawai`i

Endemic species

Percent endemism

Lineages in Hawai`i

Taxa in Hawai`i

141

141

100

1

154

Asteraceae

98

97

99

10

129

Rutaceae

56

56

100

3

58

Rubiaceae

55

53

96

11

62

Gesneriaceae

59

59

100

3

61

Cyperaceae

46

17

37

37

52

Poaceae

49

41

84

25

52

Caryophyllaceae

41

41

100

3

42

Fabaceae

26

17

65

19

26

Malvaceae

25

24

96

13

31

Piperaceae

25

23

92

4

25

Arecaceae

22

22

100

1

22

Primulaceae

20

20

100

2

20

Campanulaceae

31

Subalpine shrubland, Haleakalā, East Maui

CHAPTER 3

Plant Communities

T

must endure intense solar radiation, salinity, nutrient-poor soil, and, on dunes, burial by moving sand. Nevertheless, plants with adaptations to survive these conditions are highly successful. One component of ecological success in such coastal habitats is the ability to reproduce and colonize open habitats in this typically dynamic environment. Frequent colonization and recolonization adds to gene flow and limits the likelihood of speciation. It is not surprising, then, that many of the native species in these habitats in the Hawaiian Islands have widespread patterns of distribution across the Pacific and are indigenous rather than endemic. The common species encountered along the immediate coast in such habitats include shrubs such as Scaevola taccada (formally known as S. sericea) and Vitex rotundifolia, vines such as Ipomoea pes-caprae and Vigna marina, herbs such as Sesuvium portulacastrum, Heliotropium anomalum, and H. curassavicum, and the grass Sporobolus virginicus. In addition to these common species, there are other rare indigenous strand species such as Cressa truxillensis and Lysimachia mauritiana, whose limited distributions suggest a recent colonization of the Islands. Overall, one-quarter of all of the indigenous but nonendemic species in the Hawaiian Islands occur in coastal strand environments.

here have been many descriptive systems for classifying plant communities in the Hawaiian Islands. These tend to suffer from being much too simple to be very functional or being much too complex and difficult to assign. Here we err on the side of a descriptive system that separates seven groups of plant communities. These are coastal habitats, lowland dry forests and shrublands, mesic seasonal forests, montane wet forests, bogs, subalpine communities, and alpine communities. The most critical factor in determining landscape patterns of plant communities is the relative availability of water. This means that both elevation and slope facing are important determinants of the annual and seasonal patterns of precipitation. As described in chapter 1, rainfall gradients in the Hawaiian Islands are among the sharpest in the world because of the strong influence of orographic effects due to the steep mountains and moist trade winds. Low-lying islands in the rain shadow of higher islands experience rainfall as low as 250–300 mm, while the high volcanoes above the trade wind inversion receive little more than this. In contrast, mid elevations on the windward slopes of the higher islands have some of the highest rainfall totals anywhere in the world. A second set of factors important in determining community structure are soil factors related to the composition and age of the parent material as well as other traits of soil organic matter and drainage. On most of the islands, this parent material has formed from volcanic flows, with varying periods of weathering depending on the age of the island and the activity of individual volcanoes. Waterlogged areas of high rainfall and impeded drainage typically accumulate organic matter and form bogs. Also present are coastal areas with other forms of parent material, such as dunes and limestone outcrops. A final major factor determining community structure is temperature, with low temperatures becoming a limiting factor for many tropical species on the high volcanoes. The high volcanoes of Mauna Loa, Mauna Kea, and Haleakalā support an alpine habitat where the combined stresses of low nocturnal temperatures, drought, and high winds limit the coverage and diversity of plant species.

Coastal Habitats COASTAL STRAND AND SHORE CLIFFS Plant communities along the immediate coast on dunes and coral outcrops are impacted by a variety of stressful conditions. Plants

3.1 Makapu‘u Beach, O‘ahu

33

34

PLANT COMMUNITIES

3.2 

Coastal dunes at Mo‘omomi Reserve, Moloka‘i

Dry coastal grasslands once covered dune areas on the Northwestern Hawaiian Islands and all of the main islands, but few extensive areas of this community remain today. Examples can be seen at Kahuku on O‘ahu and Mo‘omomi on Moloka‘i. Sporobolus virginicus is often the dominant species on these dunes, along with other littoral species mentioned above. Dune grassland associations dominated by Eragrostis variabilis or Lepturus repens can be found in the Northwestern Hawaiian Islands. In addition to these indigenous strand species, there is an interesting group of endemic coastal subshrubs or shrubs that have secondarily evolved into coastal strand environments from dry upland ancestors. Examples include Euphorbia degeneri, Melanthera integrifolia, Scaevola coriacea, Phyllostegia variabilis, Kadua littoralis, and Achyranthes splendens. Two other interesting endemics in coastal strand habitats are prostrate forms of Chenopodium oahuense, a remarkably plastic shrub that can be found from the coast to subalpine shrublands, and Nama sandwicensis, an inconspicuous herb with likely ancestry from the arid Southwest of North America. Despite the widespread occurrence of many introduced trees such as sea grape (Coccoloba uvifera), kiawe (Prosopis pallida), tree heliotrope (Heliotropium argenteum), false kamani (Terminalia catappa), and coconut palms (Cocos nucifera) along Hawaiian beaches, there are no native tree species in this habitat. Coastal cliffs and lava slopes along the coast are also influenced by salt spray. These habitats are typically dominated by a mix of indigenous and endemic species, including Sida falax,

Boerhavia repens, Melanthera integrifolia, and Fimbristylis cymosa. The inner boundary of this coastal strand occurs where salt input is no longer an ecological factor and where dunes, marine terraces, and coral outcrops give way to more upland habitats.

COASTAL SHRUBLANDS AND WOODLANDS Above the strand area of the Hawaiian Islands there are distinctive coastal habitats from near sea level up to about 300 m elevation that are not influenced by salinity. Arid coastal communities occur on the leeward sides of the islands, where annual rainfall varies from 375 up to 1,000 mm and summer conditions offer extended drought. This zone has been so altered by human activities that it is unclear what the natural vegetation would have included. Three native tree species in this habitat are Erythrina sandwicensis, Santalum ellipticum, and Myoporum sandwicense. The latter species is extremely plastic in habitat distribution. Like the coastal strand, many of the common species of this zone are indigenous rather than endemic. These include Waltheria indica, Sida fallax, Caesalpinia bondoc, Plumbago zeylanica, and Osteomeles anthyllidifolia. Endemic species in this habitat include Capparis sandwiciana, Gossypium tomentosum (a tetraploid cotton that is presumably derived from a neotropical ancestor), and the rare Sesbania tomentosa. The water fern Marsilea villosa, one of the few endemic Hawaiian aquatic plants, once occurred in coastal arid habitats on Ni‘ihau, O‘ahu, and Moloka‘i, but it is known today from just

PLANT COMMUNITIES

Open dry woodlands and forests once covered large areas of the lowlands on the leeward slopes and rain shadow areas of the Hawaiian Islands, including much of the area of the smaller islands. These are generally habitats below about 1,500 m elevation that receive less than 1,300 mm of annual rainfall, with 500 mm typical of coastal areas, increasing to 750 mm at midlevel slopes. The drier western slopes may receive as little as 250 mm of rain. These forests can be open with a grassy understory or develop a closed canopy in more mesic sites with a diverse assemblage of native understory shrub. Canopy heights average about 10 m, but they can reach twice this height in favorable sites.

Lowland dry forests are generally open in structure, although they may have a closed-canopy structure at the wetter margin of their occurrence. These stands lack the tree ferns and epiphytes typical of wet forests. In their simplest form, lowland dry forests may exhibit single- or 2-species domination, with Diospyros sandwicensis as the most widespread and important species. It occurs on all of the major islands except Ni‘ihau and Kaho‘olawe. Relatively pure forest stands of Metrosideros polymorpha with a low closed canopy once occurred on lowland slopes of all the major islands, but most of this area has since been converted to agriculture. These dry forest stands can still be found on rocky lava flows with subsurface water on Maui and Hawai‘i. More commonly, however, these dry forests consist of a mix of many tree species, including local endemics. Indeed, in terms of tree species, these were once among the most biodiverse forest stands of the Islands. While most of the dry forest tree species have coriaceous evergreen leaves, there are several prominent deciduous species that lose their leaves during the dry summer months. The most common and widespread of these are Erythrina sandwicensis and Polyscias sandwicensis (formerly Reynoldsia sandwicensis), which often occur in the driest areas. There is a remnant grove of Erythrina on Kaho‘olawe, providing evidence that dry forest was once present on this island. There is also an early report from the mid-nineteenth century by Jules Remy of the presence of Neraudia on Kaho‘olawe, where it is absent today. Very little of these dry forest communities remain today in even a seminatural state. Hawaiian dry forests, like many tropical dry forests globally, have been heavily impacted by both direct and indirect human activities. No other vegetation zone on the Islands has been so significantly altered by land clearance, fire, feral goats, cattle, and invasive alien species. As a result, less than 10% of the original extent of these forests remains today. Alien woody species such as Leucaena leucocephala, Prosopis pallida, Schinus terebinthifolius, and Lantana camara and grasses such as Cenchrus setaceus, C. ciliaris, and C. clandestinus (the genus Pennisetum has now been placed within Cenchrus), Andropogon virginicus, and Schizachyrium condensatum have taken over much of the original area of dry forests (see chapter 7).

3.3 Mo‘omomi coastal scrub, Moloka‘i

3.4 

a small number of populations on the latter two islands. This fern requires periodic flooding for reproduction, followed by a decrease in water levels for the young plants to establish. It grows only in shallow depressions in clay soil or sand dunes overlaid with alluvial clay. Most Hawaiian coastal habitats are now overwhelmingly dominated by nonnative alien species. The dominants of these, as described in more detail in chapter 7, are Leucaena leucocephala, Prosopis pallida, and Casuarina equisetifolia. Leucaena often forms virtually impenetrable thickets on dry coastal slopes. Coastal areas on the windward sides of the islands experience higher levels of rainfall and less extended summer drought. Mesic coastal forests once existed in these areas, with Pandanus tectorius and Hibiscus tiliaceus as dominants, but only small areas of these forests exist today. Dry to mesic forests of Metrosideros polymorpha and Dio­ spyros sandwicensis are thought to have been present close to sea level on the windward coasts of Moloka‘i and Hawai‘i before the arrival of the Polynesians. Two Polynesian introductions of trees, Aleurites moluccana and Thespesia populnea, with the latter possibly also indigenous, are widespread today in these habitats along with a variety of alien shrubs and vines. Three other Polynesian introductions, Tacca leontopetaloides, Cordyline fruticosa (formerly called C. terminalis), and Morinda citrifolia, are also present (see chapter 6).

Lowland Dry Forests and Shrublands

Dry forest at Pa‘aiki Valley, Kaua‘i

35

36

PLANT COMMUNITIES

3.5 

Dry forest with Diospyros sandwicensis, Maui

3.7 

‘Ulupalakua dry forest, East Maui

The degradation of lowland dry forests today is such that there are only four areas with moderate-sized remnant forest patches remaining today, although individual dry forest species range more widely on dry slopes. These are stands at Mokulē‘ia on northwestern O‘ahu, at Kānepu‘u on northwest Lāna‘i, at Auwahi on the southern slopes of Haleakalā, and Pu‘u Wa‘awa‘a on the Kona slopes of Hawai‘i. Even these stands are heavily fragmented and impacted by grazing and invasive species. Dry forests at Mokulē‘ia lie on the north-facing slopes of the northwestern Wai‘anae Mountains. These forests are 10–15 m tall and range from closed-canopy forests of Diospyros sandwicensis, Metrosideros polymorpha, and Psydrax odoratum to semideciduous forest stands with Sapindus oahuensis. The small area of dry forest at Kānepu‘u on Lāna‘i lies at about 520 m elevation and is thought to be a remnant of a much more widespread forest area in the past. The dominant species here are Nestegis sandwicensis and Diospyros sandwicensis. Grazing from introduced axis deer and invasive plant species such as Lantana camara have heavily impacted this area.

3.6 

Dry forest with Erythrina sandwicensis, Kona District, Hawai‘i

Both Maui and Hawai‘i once possessed extensive areas of diverse dry forest. More than 80% of these tree species had Native Hawaiian uses for medicines, toolmaking, canoe construction, kapa making, dye production, and religious rituals. On the south slopes of Haleakalā, the highest diversity of dry forest tree species is found in the upper dry forest zone at about 900–1,500 m elevation, where annual rainfall averages 750–1,000 mm and cloud cover adds fog drip. Below 900 m the diversity drops steadily as rainfall is reduced from 750 mm down to about 250 mm. Joseph Rock visited this area in 1913 and described it in his book, Indigenous Trees of the Hawaiian Islands, as one of the most significant biological sites in all of the Hawaiian Islands. Upon his return to the area a quarter century later in 1939, Rock is said to have wept over the dramatic deterioration during his absence. This dry forest has continued to be degraded. Only an estimated 4% of the original extent remains today, but active restoration efforts are underway and showing success. Dry forests once covered most of the North Kona area on Hawai‘i with a mosaic of diverse species assemblages. The most

PLANT COMMUNITIES

3.8 

Dryland forest, Pu‘u wa‘awa‘a, Hawai‘i

Pili grassland with Heteropogon contortus. Līhau, West Maui

3.9  Dry montane shrublands at Pōhakuloa, Hawai‘i, with major invasion of alien plant species

3.10 

common native trees in this area today are Diospyros sandwicensis and Sophora chrysophylla, along with the shrubby Nototrichium sandwicense. Today, however, the dry forest ecosystem is largely represented by fragmented and depauperate patches of native trees, scattered in a matrix of the alien fountain grass Cenchrus setaceus. This invasive grass, discussed in more detail in chapter 7, outcompetes and suppresses reproduction of native woody species and promotes frequent burning, to which these native species are poorly adapted. On Hawai‘i, dry forests grade into dry shrublands at higher elevations in the saddle area between Mauna Kea and Mauna Loa. Eight tree species are widely present today in relict dry forest areas. These are Diospyros sandwicensis, Erythrina sandwicensis, Metrosideros polymorpha, Myoporum sandwicense, Nestegis sandwicensis, Psydrax odoratum, Polyscias sandwicensis, and Xylosma hawaiiense. Many other common species from these sites are restricted to a single island. Shrublands replace dry forests on many of the driest lowland slopes, particularly in areas with rocky substrate and hot,

dry summers. One of the most widespread of these communities is a low shrubland of Dodonaea viscosa, which can be found on all of the major islands. A variety of native shrub species may be codominant in these stands. Open shrublands dominated by native species of Bidens can be found on exposed lava cliffs of the Wai‘anae Mountains on O‘ahu and on Moloka‘i and West Maui. Open shrublands of Wikstroemia species are widespread at low elevations on windward slopes of Moloka‘i, Maui, and Hawai‘i. Common associated species in all of these shrublands include Euphorbia celastroides, Myoporum sandwicense, and Osteomeles anthyllidifolia. While invasive alien grasses cover much of the open lowlands of the Hawaiian Islands, there are lowland grasslands dominated by a native species. These are the pili grasslands of Heteropogon contortus, on lowlands cleared and maintained by the Native Hawaiians with frequent fires. The pili grass was widely used in thatching. These communities are not stable with time, however, and are eventually colonized by woody plants.

37

38

PLANT COMMUNITIES

Mesic Seasonal Forests Mesic seasonal forests once covered broad areas of the Hawaiian Islands, lying on the windward slopes both below and above wet forest and on the leeward slopes above dry forests. On Hawai‘i, however, these forests reappear above the montane wet forests because of the limiting moisture conditions produced by the trade wind inversion. Summers are dry, but annual rainfall levels range from 1,200 to 3,800 mm, with the majority of this precipitation falling from October to March. The high levels of rainfall compensate for the drier summer conditions that affect the dry forest communities more strongly. Mesic forests can take a variety of associations with variations in dominant species. Rainfall, substrate age, soil drainage, and soil nutrient availability are important factors determining species composition. The height of forest canopies, tree density, and diversity of tree species on any single island are generally positively correlated with elevation, rainfall, and soil fertility. The most diverse lowland mesic forests occur on the older islands of Kaua‘i and O‘ahu, but taller forests occur on Maui and Hawai‘i.

3.11 Mesic ‘ōhi‘a forest, Hawai‘i Volcanoes National Park, Hawai‘i

Two widespread mesic forest types can be recognized: ‘ōhi‘a forest and koa forest, as well as a series of more locally distributed forest communities. Lowland mesic forests of ‘ōhi‘a occur on all of the major islands, with the exception of the more arid Ni‘ihau and Kaho‘olawe. The nearly ubiquitous Metrosideros polymorpha is the dominant species in these forests. The composition of associated canopy and subcanopy tree species varies widely over the range of this community. Some ‘ōhi‘a forests have Psychotria mariniana (Wai‘anae Mountains on O‘ahu) or P. hawaiiensis (South Kona District, Hawai‘i) as codominant species. There are long lists of other tree species that may be present. An interesting form of mesic ‘ōhi‘a forest grows on the south-facing slopes of Kīlauea at 400–1,200 m elevation on young lava flows less than 1,000 years in age. These are pioneer forest communities, representing an early stage of succession. The open tree cover in this area is overwhelmingly dominated by Metrosideros polymorpha, with scattered shrubs of Dodonaea viscosa and Leptecophylla tameiameiae (formerly called Styphelia tameiameiae) in the openings between trees. This community has been heavily invaded over the past four decades by several alien species that have dramatically altered the ecosystem

PLANT COMMUNITIES

3.12 

Mesic diverse forest with mixed dominance of koa and Aleurities moluccana, Ko‘olau Mountains, O‘ahu

structure and function. Morella faya (formerly called Myrica faya), a nitrogen-fixing species from volcanic substrates in the Canary Islands, was an early invader that could become established on this young substrate. Two grass species, Andropogon virginicus and, more significantly, Schizachyrium condensatum, have spread as virtual ground covers, aided in part by the increased nitrogen produced by Morella. As discussed in chapter 7, these grasses have dramatically increased the frequency and intensity of fires in this habitat, to the great detriment of many native species that do not readily resprout after burning. Forests dominated by Acacia koa occur on all of the six larger islands. On the older islands of Kaua‘i and O‘ahu, these forests generally lie below the montane forests of Metrosideros, on sites less moist and better drained than those that support wet forest. These are generally highly diverse forests. Metrosideros polymorpha is often present, along with other tree species such as Bobea elatior, Charpentiera obovata, Myrsine kauaiensis, Nestegis sandwicensis, Psychotria kaduana, P. mariniana, Pisonia umbellifera, Psydrax odoratum, Santalum freycinetianum, and Sapindus oahuensis. Common understory shrubs include Pipturus albidus, Scaevola chamissoniana, S. gaudichaudiana, and Wikstroemia oahuensis, as well as the seemingly ubiquitous Dodonaea viscosa and Leptecophylla tameiameiae. Unlike the higher rainfall wet forest above, epiphytes are few in these stands. Montane mesic forests of Metrosideros polymorpha and Acacia koa occur at middle elevations up to 2,000 m on East Maui and the higher volcanoes on Hawai‘i and to a lesser extent on Kaua‘i and West Maui. This is a zone above the wet forest belt, with a seasonal rainfall regime and 1,000–1,900 mm of annual rainfall. Soils are highly varied, ranging from young lava flows to soils that are weathered sufficiently to provide fertile conditions for tree growth. Canopy ranges from moderate heights no more than 10 m on shallow soils to forests up to 35 m tall on deep soils of volcanic ash. This community can take several forms. On the leeward slopes of East Maui and Hawai‘i, it may be strongly dominated by Metrosideros with a closed canopy 10–25 m in height and

3.13 

Wet koa forest, Makawao Forest Reserve, East Maui

well-developed layers of subcanopy trees and understory shrubs. Unlike the wet forests of Metrosideros on Hawai‘i, Cibotium tree ferns are not important community components. A taller community on these two islands, as well as at about 1,000 m elevation near Kōke‘e on Kaua‘i, has a mixed dominance of Metrosideros and Acacia koa, with the latter forming emergent trees 35 m in height. A third codominant canopy tree, Sapindus saponaria, is present in mixed koa and ‘ōhi‘a forest in Hawai‘i Volcanoes National Park. Open koa forests on relatively young lava and ash flows at these elevations are relatively low in species diversity, but they are not pioneer communities like those of the lower-elevation Metrosideros stands on young lava flows. A common associated tree is Metrosideros polymorpha var. incana. Also present are Sophora chrysophylla and Myoporum sandwicense, small trees that form subalpine woodlands at higher elevations. Arid adapted shrubs such as Dodonaea viscosa, Leptecophylla tameiameiae, Vaccinium reticulatum, and the native grass Deschampsia nubigena occur among the trees. Large areas that once supported ‘ōhi‘a and koa/‘ōhi‘a forest have been logged for the koa and converted to cattle ranching. Once this conversion takes place, the establishment of alien pasture grasses such as Cenchrus clandestinus and Melinis minutiflora makes tree restoration very difficult. Feral goats and pigs have damaged much of the remaining native stands. Distinctive local stands of mesic forest communities often occur in moist gulches and stream canyons at the head of steep valleys. On Kaua‘i and O‘ahu, Pritchardia species may form pure stands in such sites, with the palms reaching heights up to 10–12 m. More widespread are riparian communities of Pisonia and Charpentiera in shaded canyons on all of the larger islands. Perhaps the easiest to recognize of such steep valley communities, however, are ones dominated by Aleurites moluccana, a Polynesian introduction. The pale green foliage of this tree stands out from the color of other leaves, allowing it to be recognized from a great distance.

39

40

PLANT COMMUNITIES

3.14 Wet ‘ōhi‘a forest, Pu‘ukukui, West Maui

Montane Wet Forests Montane wet forests are sometimes termed montane rain forests or cloud forests in the Hawaiian Islands. These forest areas generally occur at elevations of about 1,000–1,900 m, which mark the lower and upper boundaries of the cloud zone as produced by the trade wind inversion. The moist cloud cover that reaches to ground level in this zone provides cool growing conditions with high rainfall and abundant fog drip. Typical annual rainfall levels in these habitats are extremely high, exceeding 2,500 mm and reaching as high as 7,000 mm or more on Kaua‘i and Maui. These low forests typically present a dense tangle of lichen and moss-covered branches and an understory with rotting logs. Vascular epiphytes, particularly ferns, grow luxuriantly in the foggy mist that regularly bathes these areas. On the older and lower islands, wet forests are present in areas along upper ridges and crests facing the trade winds, where cloud layers form or saturate the area for much of the day. Often these forests are patchy and fragmented because of the topography. The best examples of these forests on the older islands can be seen in the northern Alaka‘i Swamp of Kaua‘i and the area around Mount Ka‘ala on O‘ahu, where they occur on relatively level plateaus. Similar cloud forests occur on the windward slopes of higher islands, as with the Waikamoi forest at about 1,200–1,900 m on the west slope of Haleakalā and the Kīlauea forest at 1,500–1,900 m on the wet east-facing slopes of Mauna Loa. Smaller elements of cloud forest communities can be found on the high windward ridges of Moloka‘i as well.

3.15 

Diverse wet forest at Pu‘ukolekole, Moloka‘i

PLANT COMMUNITIES

3.16 

Cibotium understory in wet ‘ōhi‘a forest, Hawai‘i

Tree architecture is varied in Hawaiian wet forests, with gradients in the growth form and height of forest trees related to substrate age and drainage as the major factors defining this trait. The heavily weathered areas of Kaua‘i and O‘ahu with wet forests exhibit boggy soils, as the high rainfall and relatively level topography produces poor drainage, with deep mucky soils over gray clay substrate indicative of anaerobic conditions. Such conditions lead to a low stature and gnarled forest trees, typically only 4–7 m in height. Better soil drainage in cloud forests on the west slopes of Haleakalā allows better growing conditions, and forests reach 8–15 and sometimes 20 m in height. Finally, the relatively young, wet forest soils on the Hawaiian Islands have fertile and well-drained substrates that allow forests to reach up to 30 m in height. Several tree species are common and often codominant in wet forest areas of Kaua‘i, O‘ahu, Maui, and Hawai‘i. These are a glabrous form of ‘ōhi‘a (Metrosideros polymorpha var. glaberrima), Acacia koa, and Cheirodendron trigynum. Also common in all four wet forests are Myrsine lessertiana, Ilex anomala, and Kadua affinis (formerly Hedyotis terminalis). There are interesting additional patterns of speciation, however. A pubescent variety of ‘ōhi‘a, M. polymorpha var. polymorpha, is present on three of the islands but absent from Kaua‘i. Two additional species of Cheirodendron have limited ranges, with C. faurei restricted to Kaua‘i and C. platyphyllum on both Kaua‘i and O‘ahu. Large tree ferns are often important components of the canopy or subcanopy on windward areas of Hawai‘i, with Cibotium glaucum often forming a continuous subcanopy and C. chamissoi and

C. menziesii also present. The dense understory of C. glaucum and Sadleria pallida once present in wet forests on Maui has been heavily impacted in the past by feral pigs, cattle, and goats. Two families, the Rubiaceae and Rutaceae, show strong patterns of speciation among wet forest trees, with species replacements in similar ecological habitats as one moves between islands. In the Rubiaceae, for example, Kaua‘i has 3 common and endemic species of wet forest Coprosma—C. elliptica, C. kauensis, and C. waimae—but lacks the widespread C. ochracea; O‘ahu has only C. ochracea and the endemic C. longifolia in mesic forests. Moloka‘i and West Maui share the endemic C. ternata, while East Maui and Hawai‘i share C. rhyncocarpa in this habitat. A similar pattern of local wet forest endemics continues with a speciose group of Rutaceae. Melicope feddei and M. cruciata, among several other species of this genus, occur only on Kaua‘i, while M. christophersenii and M. oahuense are examples of wet forest species restricted to O‘ahu. This pattern continues with M. haleakalae and M. orbicularis among species endemic to Maui. Hawai‘i has just 2 endemic taxa of the speciose genus: M. radiata and the endangered M. zahlbruckneri. While Joseph Rock celebrated the diversity of the Hawaiian dry forest trees, wet forests in the Hawaiian Islands are far richer in species when we add woody shrubs, ferns, and herbs to the community floras. As with the trees, there is a notable pattern of speciation in many shrub groups. Wet forest habitats are home to a diverse assemblage of species of Cyrtandra, Myrsine, Stenogyne, Labordia, Clermontia, Cyanea, and Dubautia. The moist conditions in these wet forests make them excellent habitats for more

41

42

PLANT COMMUNITIES

3.17 

Diverse wet forest, Kohala Mountains, Hawai‘i

than 60 species of ferns, with Asplenium, Adenophorus, Elaphoglossum, Grammitis, Cibotium, and Dryopteris as diverse groups with 3 or more wet forest species. Touchardia latifolia represents a monotypic endemic genus of shrubby wet forest trees present on all of the major islands. This species was important for the Native Hawaiians as a source of fiber for rope. Three wet forest species of Euphorbia (formerly separated as Chamaesyce) are broad-leaved shrubs evolved from small-leaved ancestors from drier habitats. These are E. clusifolia and E. rockii on O‘ahu and E. remyi on Kaua‘i. Two species of Gunnera, unusual large-leaved herbs, can be found on cloud-swept wet forest ridges. Gunnera kauaiensis is restricted to Kaua‘i, while G. petaloidea is found at Mount Ka‘ala on O‘ahu and on eastern Moloka‘i, Maui, and the Kohala Mountains of Hawai‘i. Like other Gunnera species, these plants have symbiotic colonies of nitrogen-fixing cyanobacteria within the tissues of their rhizomes. Extensive thickets of Dicranopteris linearis, an indigenous fern common in the Old World tropics and Polynesia, occur widely in Hawaiian wet forests and mesic forests. This fern colonizes secondary successional sites of landslides, roadcuts, and other disturbed areas, and once established it may persist for many years. Tree succession, often very slow because of the density of the Dicranopteris canopy, eventually shades out these thickets.

Bogs Although the high rainfall in montane areas of the five larger islands would suggest that bogs should be common, in reality Hawaiian bogs are relatively small in number and size. Wet bogs occur in scattered locations of poor drainage in the montane wet forests of the five largest islands. The most prominent bogs occur in the Alaka‘i Swamp of Kaua‘i. The name and broad extent of the Alaka‘i Swamp suggest a large area of bog habitat, but in reality the name “swamp” is misleading. The great majority of the area is low Metrosideros wet forest, with scattered small bogs across the plateau.

3.18 Alaka‘i Swamp trail, Kaua‘i

Other prominent bogs are on Mount Ka‘ala on O‘ahu, Pēpē‘ōpae Bog on East Moloka‘i, Pu‘ukukui and Mount ‘Eke on West Maui, smaller bogs on the northeast slope of Haleakalā, and the summit area bog of the Kohala Mountains on Hawai‘i. The general elevational range of bogs in the Hawaiian Islands is about 1,100–1,700 m, corresponding to the zone of maximum rainfall from the trade wind inversion. However, bogs can occasionally occur at lower or higher elevations. Wahiawa Bog, in a rugged area of the south of Kaua‘i, occurs at only 630 m, while the uppermost bog on Haleakalā lies at 2,270 m in an area of subalpine grassland. Generally bogs occur on flat or gently sloping surface with impervious clay substrates that prevent water from draining into the soil. With impeded drainage, boggy conditions will form where rainfall exceeds evapotranspiration. The impervious layer often results as humic acid from decomposing vegetation helps to break down volcanic basalts and form fine-textured clays. Once drainage becomes greatly impaired, peats begin to form, with substantial accumulation as deep as several meters, underlain by gray anaerobic clays. These conditions of waterlogging make it difficult for tree species to grow roots and survive, producing conditions supporting low sedge bogs or shrublands. An exception to the general pattern of bogs occurring on flat areas can be found with bog areas on relatively steep slopes near the summit region of Wai‘ale‘ale on Kaua‘i. The immense quantity of rainfall here keeps soil permanently saturated with water. The flora of Hawaiian bogs is an interesting one in that in many floristic respects, it is more typical of the southern hemisphere than the northern hemisphere. At the generic level, there are strong affinities with bog plants from Malaysia but also with temperate bogs in New Zealand and southern Chile. Many of the characteristic bog species are not entirely restricted to this habitat and can often be found in open areas of wet montane forest or at the upper margin of these forests in the transition to subalpine shrublands. The classic open bogs on Kaua‘i, Moloka‘i, Maui, and Hawai‘i are dominated by irregular low hummocks of cushion-form sedges, with grasses and low shrubs contributing smaller

3.19 Pēpē‘ōpae Bog, Moloka‘i

3.21 

Keahikauō Bog, West Maui

3.20 Mount ‘Eke Bog with Argyroxiphium calignis, West Maui

44

PLANT COMMUNITIES

3.22 

Wahiawa Bog, Kaua‘i

3.23 

Greensword Bog with Oreobolus, East Maui

amounts of cover. Larger shrubs and trees are generally restricted to the bog margin where drainage is better. The dominant species is commonly the cushion-forming sedge Oreobolus furcatus, with 2 other sedge species, Rhynchospora rugosa subsp. lavarum and R. chinensis subsp. spiciformis, taking on similar growth forms. Carex echinata is a dominant species in East Maui bogs. Large tussocks are formed by the bog grasses Dichanthelium cynodon and D. hillebrandianum. A trailing gray moss, Rhacomitrium lanuginosum, is common and often forms a major component of cover. While Sphagnum would be expected to form a significant component of such boggy habitats, this moss genus is restricted in distribution to bog areas of the Kohala Mountains and Mount Ka‘ala, with this latter occurrence thought to be due to a deliberate introduction by Charles Forbes. Grasses, herbs, and ferns are also present in the bogs, along with scattered woody shrubs. The most common tussock-forming

3.24 

Greenswords in Mid Camp Bog, East Maui

grasses are species of Dichanthelium and Deschampsia nubigena. In many cases, the bog shrubs are dwarfed forms of species that grow larger in other habitats. Good examples of this dwarfing can be seen with Metrosideros, Cheirodendron, and Vaccinium species, the common canopy trees and taller shrubs in surrounding areas of wet forest. Dwarf Metrosideros in bogs include 3 distinct shrubby taxa: M. polymorpha var. pumila (Kaua‘i, O‘ahu, and Moloka‘i), M. polymorpha var. pseudorugosa (West Maui), and M. waialealae (Kaua‘i, Moloka‘i, and Lāna‘i). The Hawaiian tarweed group has evolved a number of interesting bog species. The most iconic of these are 3 bog species of Argyroxiphium, close relatives of the silversword. Two of these are found in Maui bogs. These are the ‘Eke silversword, A. caliginis, on ‘Eke and Pu‘ukukui on West Maui and the greensword, A. grayanum, found on bog margins on West Maui and bogs on East Maui. The Ka‘ū silversword, A. kauense, is limited in distribution to two small areas of forest openings and bogs on Hawai‘i

PLANT COMMUNITIES

at elevations of 1,600 to 2,320 m, with the total population close to extinction before recent conservation efforts to protect it from feral grazing animals and pigs. There are 5 bog-edge species of Dubautia, with D. imbricata, D. paleata, D. raillardioides, and D. waialaeae endemic to Kaua‘i and D. laxa occurring from Kaua‘i to Maui. Four species of Hawaiian Lobelia are bog specialists, with L. kauaiensis and L. villosa on Kaua‘i, L. gaudichaudi in the Ko‘olau Mountains of O‘ahu, and L. gloria-montis on Pu‘ukukui on West Maui and the East Maui bogs. A number of herbaceous bog species are characterized by a rhizomatous growth form with multiple points of rooting across the bog surface. This form can be seen in bog species of Keysseria, Viola, and Astelia and similarly in the trailing stems of Argyroxiphium caliginis and Geranium hillebrandii in the bogs on West Maui. An unusual bog specialist is Drosera anglica, a widespread carnivorous plant more characteristic of north temperate bogs. It occurs only on Kaua‘i.

Subalpine Communities Subalpine and alpine environments are separated from the lower forest areas by two significant climate boundaries that occur with increasing elevation. The first of these is the upper level of the cloud zone, which commonly occurs at about 1,800–2,000 m. Because the moist trade winds reach an inversion layer at this elevation, rainfall peaks in the upper montane wet forests below, separating forest vegetation from drier subalpine woodlands and shrublands. Rainfall declines steadily from about 1,300 mm annually at the lower margin of the subalpine zone to 500 mm or less at the highest alpine elevations of Mauna Loa and Mauna Kea. The second climate boundary is that representing regular frost conditions, which has been used to separate subalpine from alpine zones. Having described these boundaries, it is perhaps fair to state that these are more conceptual than real, as

3.25 

Subalpine shrubland, Haleakalā, East Maui

3.26 

they do not plot readily with elevation and slope, and they show distinctive features on each of the high volcanoes of Mau‘i and Hawai‘i. Plants in the upper subalpine and alpine environments of the Hawaiian Islands frequently experience subfreezing temperatures during the night, especially during the winter months. Subalpine shrublands and high-elevation woodlands encircle the upper slopes on both the windward and leeward slopes of Haleakalā, Mauna Loa, Mauna Kea, and Hualālai. Within these belts, the composition and structure of plant communities vary with elevation, rainfall, and substrate age. The lower subalpine zone on Haleakalā and scattered areas among the Sophora/Myoporum woodland on Hawai‘i support a shrubland with mixed dominance of species. In general appearance and structure, this community is much like California chaparral, with a moderately continuous cover of evergreen sclerophyllous shrubs 1.5–3.0 m in height. This community is commonly termed “styphelia scrub” because of the major coverage by Leptecophylla tameiameiae. On Haleakalā, the codominant species in these shrublands include Coprosma montana, Dubautia menziesii, Geranium cuneatum, Vaccinium reticulatum, Sophora chrysophylla, and the prostrate Coprosma ernodeoides in open areas. Santalum haleakalae var. haleakalae is a common endemic in this community on Haleakalā. Open areas among the shrubs also support bracken fern, Pteridium aquilinum, and a variety of temperate-climate native grasses such as Deschampsia nubigena and Trisetum glomeratum. The high-elevation slopes on Hawai‘i are dominated over broad areas by a woodland of Sophora chrysophylla and Myoporum sandwicensis. Common shrub associates are Leptecophylla tameiameiae, Vaccinium reticulatum, and Chenopodium oahuense. Typical examples of this woodland can be seen on Mauna Kea from about 1,900 to 2,500 m elevation. At its lower margin here, this community takes the form of an open forest with scattered clumps of Acacia koa. On Mauna Loa, this community is best

Dry subalpine woodland on Mauna Kea, Hawai‘i

45

46

PLANT COMMUNITIES

developed at elevations of 2,000–2,500 m, with Sophora and Metrosideros often sharing dominance and Myoporum less abundant. The presence of Metrosideros here, in contrast to its absence from this zone on Mauna Kea, is thought to represent a successional development on younger soils. The relatively low treeline of Metrosideros on Mauna Loa compared to treelines of other species in continental areas is thought to be due to the susceptibility of local populations of Metrosideros to freezing damage at relatively mild frost conditions. Other physiological ecotypes of Metrosideros have been shown to be less susceptible to frost damage. In contrast to Mauna Loa and Mauna Kea, the windward slopes of Haleakalā lack Sophora/Myoporum woodland, although Sophora is present. Native Hawaiians had little impact on the structure of Sophora/Myoporum woodland because of its elevation. However, grazing by feral cattle, sheep, goats, and pigs have all had major effects on subalpine communities over the past two centuries. These impacts are most prominent on Mauna Kea, where grazing has been very pronounced. Here, various alien grasses, most notably Holcus lanatus, have heavily invaded at the expense of native grasses. In some areas, native shrubs such as Dodonaea viscosa and Wikstroemia sandwicensis that are relatively unpalatable to livestock have expanded their dominance at the expense of other native species. Local areas in the lower subalpine zone on Hualālai and Mauna Kea support an unusual shrubland dominated by Chenopodium oahuense. For a temperate-region botanist, it seems odd to think about Chenopodium as a woody plant. Nevertheless, C. oahuense has evolved as a shrub or even a small tree, reaching heights of up to 4.5 m on Mauna Kea. This species is highly plastic in its ecology, being found across the full range of dryland habitats in

3.27 

Deschampsia grassland, Haleakalā, East Maui

the Hawaiian Islands, from coastal to subalpine. It does well as a colonizer on old lava flows following site disturbance. There is pollen core evidence suggesting that C. oahuense had a much more expanded range during dry interglacial periods of the Pleistocene. A fourth subalpine community present on the upper slopes of Haleakalā and Hawai‘i is a tussock grassland composed of a mixed assemblage of native bunchgrasses that reach up to 1 m in height. Large individual tussocks of Deschampsia nubigena, up to 1 m across, once dominated these high-elevation grasslands. Other perennial bunchgrasses present include C3 species such as Poa sandvicensis, Trisetum glomeratum, and Agrostis sandwicensis adapted to cool temperate conditions. Panicum tenuifolium and Eragrostis atropioides, occurring here at their upper elevational limits, are among the only warm subtropical C 4 grass species present. Other associated native species include the sedges Luzula hawaiiensis, Uncinia brevicaulis, and Carex macloviana, as well as scattered individuals of Vaccinium reticulatum and Pteridium aquilinum. Extensive rooting by feral pigs in the wetter margins of this community has reduced species diversity and facilitated invasion by the alien Holcus lanatus.

The Alpine Zone Higher-elevation areas of Haleakalā above 2,000 m elevation within the cinder-dominated crater and above 2,600 m on the older outside western slope of the volcano are often considered to represent an alpine community, although these do not possess true alpine climate regimes in the classic sense. In the crater, the alpine zone occurs adjacent to and is relatively sharply defined from relatively subalpine shrublands on older and more developed substrates. The alpine zone landscape consists of rough broken lava features, cinder cones, and cinder fields, with little soil development. Conditions within this alpine zone produce severe constraints on plant survival and growth. Stress comes both from edaphic conditions due to the young volcanic substrates and climatic conditions due to strong patterns of diurnal temperature change, frosts, and high solar radiation. As barren as this landscape appears, however, rainfall is surprisingly high, averaging about 1,000 mm annually. The alpine zone on Haleakalā is sparsely vegetated, with plant cover typically less than 5% and plant species diversity low. The most famous and charismatic species of this zone is Argyroxiphium sandwicense subsp. macrocephalum, the silversword, which is abundant across the cinder fields of Haleakalā at elevations from 2,125 m to near the summit at 3,055 m. These rosette plants grow for 20–100 years in the field before forming a massive inflorescence, flowering, and then dying. The shiny silver leaf surface and parabolic form of the silversword rosette provide warm growing temperatures above cool alpine conditions for the apical meristem at the center of the rosette. Concern about the protection of this species was a major factor in the decision to place Haleakalā within the management of the National Park Service in 1916, first as a section of Hawai‘i Volcanoes National Park and later, in 1961, as Haleakalā National Park. In addition to the large populations of silversword on Haleakalā, Argyroxiphium sandwicense subsp. sandwicense survives today in small numbers at about 2,850 m elevation in the Wailuku Valley drainage on Mauna Kea. There is evidence, however, that tens of thousands of silverswords once covered the upper slopes

PLANT COMMUNITIES

3.28 

Silversword Reserve, Mauna Kea, Hawai‘i

of Mauna Kea over elevations of about 2,600 to 3,800 m. These silversword populations began to decline in the late nineteenth century, when sheep and goats were introduced to Hawai‘i, and by the middle of the twentieth century these plants were almost entirely decimated. Two close relatives of silversword often extend up into this zone from subalpine shrublands below. These are Dubautia menziesii on Haleakalā and D. arborea on Mauna Kea. Silene struthioloides is also common in this zone. An exception to the North American origin of most Hawaiian alpine species can be seen with Tetramolopium humile, a common dwarf shrub with ancestors in New Guinea. The grasses Trisetum glomeratum and Agrostis sandwicensis are widely present but lack extensive ground cover. Dry alpine barrens are present above about 2,500–2,900 m on the high volcanoes of Mauna Loa and Mauna Kea on Hawai‘i and on the high-elevation areas of Haleakalā. Rainfall at this elevation is lower than that of the wet forests below, averaging 750– 1,250 mm annually. Frosts can occur at any time of the year, and snow may occur in winter above 3,500 m on Hawai‘i. These cold conditions, relatively low rainfall, porous and rocky soils, and intense solar radiation make this a challenging environment for plant growth. Plant cover is very low in these habitats, which take

on an appearance of a barren, lunar wasteland. It is perhaps not surprising because of the temperate nature of the environment in these alpine shrublands that many of the characteristic plant species have evolved from colonists from North America rather than from Southeast Asia and the western Pacific. The upper elevational limits of trees and thus the lower boundary of alpine habitats occurs at about 2,500 m on Mauna Loa and is marked by bushy trees of Metrosideros polymorpha 4–5 m in height. Treeline occurs 400 m higher on Mauna Kea, where Sophora reaches 2,900 m. Scattered scrubby growth of Leptecophylla tameiameiae forms the dominant plant cover above this treeline elevation. Scattered low woody plants in this zone also include Vaccinium reticulatum, Dodonaea viscosa, and Coprosma ernodeoides. There is almost no vascular plant life above 3,300 m, and only a few hardy grasses and ferns and scattered clumps of Leptecophylla tameiameiae occur above 3,000 m. The highest occurrence of any native vascular plant on Hawai‘i is at 3,970 m on Mauna Kea, where the grass Trisetum glomeratus occurs at the margin of a small remnant lake. The summits of both Mauna Loa, which is still volcanically active, and Mauna Kea present young substrates and harsh alpine conditions that are inhospitable to plant growth.

47

Lobelia kauaensis 48

CHAPTER 4

Flowering Plants

I

to standardize their source to a single reference. Habitat descriptions and elevational ranges of species are largely taken from the Manual of the Flowering Plants of Hawai‘i (Wagner, Herbst, and Sohmer, 1990; hereafter “the Manual”). The scientific names used at the family, genus, and species level also generally follow those used in the Manual, but they are updated to show subsequent changes in the current acceptance for names changed since the publication of the Manual, as shown in the Smithsonian’s online checklist, available at http://botany.si.edu/ pacificislandbiodiversity/hawaiianflora/index.htm. Changes in names from those used in the 1990 edition of the Manual are noted in the text. Species not native to the Hawaiian Islands but with cultural significance and introduced by the Polynesians before European contact are described and illustrated in chapter 6. We have selected the most ecologically significant and aggressive alien species, which are described and illustrated in chapter 7.

n this chapter, families are presented alphabetically and genera and species also alphabetically within their taxonomic group. The family names may be easily recognized, as they all end with the suffix -aceae. The scientific name of each genus and species is followed by the Hawaiian name and an indication of the species as indigenous (i.e., occurs naturally in the Hawaiian Islands as well as elsewhere) or endemic (i.e., occurs only in the Hawaiian Islands). The conservation status of species as secure, rare, vulnerable, threatened, endangered, or extinct is also shown (see chapter 8 and the appendix for definitions). “Secure” is used in the sense of not falling into one of the other categories and does not necessarily indicate the presence of a species with large and/or numerous populations. Family, genus, and species descriptions include key morphological features, habitat type of occurrence, elevational range, and geographic range of occurrence on the Hawaiian Islands and beyond. Figures on the global number of species for individual families and genera generally follow those in Mabberley (2008)

Dicotyledons

Sesuvium portulacastrum (L.) L.

Aizoaceae

INDIGENOUS | SECURE

‘ākulikuli, sea purslane

(FIG-MARIGOLD FAMILY)

Mostly succulent annual or perennial herbs, rarely low shrubs. Key features are fleshy leaves, a 5–8 lobed calyx, many stamens, and often many staminodes that are similar to petals in appearance. There are 127 genera and about 1,860 species that occur in the tropics and subtropics, mostly in South Africa and to a lesser extent in Australia. In the Hawaiian Islands represented by the indigenous genus Sesuvium. The genera Tetragonia and Trianthema are naturalized.

4.1 

Sesuvium—A genus of approximately 22 halophytic species of succulent annual or perennial herbs or subshrubs. Widely distributed in the tropics and subtropics, especially in coastal habitats. The name is in reference to the land of Sesuvii, a Gallic tribe.

Decumbent to ascending or rarely erect pantropical perennial with fleshy stems and leaves, often becoming yellow or red with age. Widely distributed along coastal sites on Midway Atoll, Pearl and Hermes Atoll, Lisianski, Laysan, Necker, and on the main Hawaiian Islands.

Sesuvium portulacastrum

49

Amaranthaceae

(AMARANTH FAMILY)

Primarily herbs, but occasionally subshrubs and rarely trees. Key features are the presence of dry, membranous bracts subtending the flowers and a usually dense or congested inflorescence. There are no petals, only a perianth comprised of sepals that may be distinct or joined at the base. There are about 175 genera and 2,000 species that are widespread in tropical and subtropical regions and a few in temperate areas. In the Hawaiian Islands represented by the endemic genus Nototrichium and the indigenous genera Achyranthes, Amaranthus, Charpentiera, and Chenopodium. Alternanthera and Gomphrena are introduced. The Chenopodiaceae, once considered a separate family, has now been merged into the Amaranthaceae.

Achyranthes—A genus of 8–10 species of mostly paleotropical annual or perennial

50

FLOWERING PLANTS

herbs or shrubs. The name is derived from the Greek achyron, chaff, and anthos, flower, in reference to the chaffy bractlike perianths. Asian species have been used in traditional Chinese medicine to treat arthritis and acute edema. In the Hawaiian Islands there are 3 endemic species representing a single colonizing lineage. Two of the species are extinct and the other is endangered.

Amaranthus brownii Christoph. & Caum ENDEMIC | ENDANGERED

Kaua‘i (Ho‘olulu, Hanakāpī‘ai, Hanakoa valleys). Known to hybridize with C. elliptica where they occur together.

Charpentiera elliptica (Hillebr.) A. Heller pāpala

ENDEMIC | SECURE

Achyranthes splendens Mart. ex Moq. ‘ahinahina

ENDEMIC | RARE/ENDANGERED

4.3 

Amaranthus brownii

Perennials with erect, densely leafy stems to 1 m long and linear, somewhat folded leaves. Found only on Nīhoa Island. Less than a dozen plants were observed in 1984.

4.2 

Achyranthes splendens var. splendens

Shrubs to 2 m tall with densely whitehairy obovate to broadly elliptic leaves. Scattered on rocky or talus slopes in open, dry forest remnants, generally from sea level to 30 m elevation. There are 2 varie­ ties, with var. spendens occurring on Lāna‘i and Maui, while the var. rotundata is an endangered shorter shrub with suborbicular leaves once found along the western coasts of O‘ahu, Moloka‘i, and Lāna‘i but now extinct on the latter two islands.

Amaranthus—A genus of about 70 species of annual or perennial herbs found in warm temperate and subtropical regions of the world. The name is derived from the Greek amarantos, unfading, in reference to the persisting color of the inflorescence. Many species are troublesome weeds, a few produce edible foliage and grains, and some have horticultural merit. In the Hawaiian Islands there are 5 naturalized and 1 endemic species.

Charpentiera—A genus of trees and woody shrubs. Of the 6 recognized species, 5 are endemic to the Hawaiian Islands and the other is known from the Austral and Cook Islands. The name honors Jean G. F. de Charpentier, a German-born Swiss naturalist and geologist. The dry wood of members of this genus is extremely lightweight and flammable.

Charpentiera densiflora Sohmer pāpala

4.5 

Charpentiera elliptica

Slender to shrubby trees to 6 m in height with somewhat coriaceous, elliptical to lanceolate leaves. Common in diverse mesic forest on Kaua‘i, particularly the Kōke‘e Plateau, at elevations of 250–1,250 m. The dry wood of this and other species is lightweight and very flammable. Used by the Native Hawaiians for pyrotechnical displays.

Charpentiera obovata Gaudich. pāpala

ENDEMIC | SECURE

ENDEMIC | VULNERABLE

4.4 

Charpentiera densiflora

Trees to 12 m in height. Leaves coriaceous, elliptic to subovate, up to 40 cm in length. Internodes between individual flowers typically only 1–2 mm. Occurs in moist valleys of Pandanus forest at 160– 250 m elevation along the Nāpali Coast,

4.6 

Charpentiera obovata

FLOWERING PLANTS

Trees to 4.5 m tall, often shrubby with obovate to somewhat elliptical leaves. Occurs in mesic to wet forest at elevations of 190–1,750 m on all of the main islands except Ni‘ihau and Kaho‘olawe; particularly common in the Wai‘anae Mountains of O‘ahu.

Chenopodium oahuense (Meyen) Aellen ‘ahea, ‘āheahea, āweoweo, goosefoot ENDEMIC | SECURE

Both Nototrichium and Charpentiera have an unusual growth form for this family as they are small trees.

Nototrichium humile Hillebr. kulu‘ī

ENDEMIC | ENDANGERED

Charpentiera tomentosa Sohmer pāpala

ENDEMIC | SECURE

4.8 

Chenopodium oahuense—coastal form 4.10 

Nototrichium humile

Erect or decumbent shrubs 1–5 m tall, openly branched. Leaves ovate to oblong 3–9 cm in length. Flowers in slender spikes 3–14 cm in length. Rare in open, remnant dry forest or in gulches, steep slopes, or cliff faces in the Wai‘anae Mountains, O‘ahu, at elevations of 300– 700 m and rare on East Maui (Luala‘ilua Hills, extinct?). There are 12 extant populations on O‘ahu with more than 3,000 individuals. Fire is the major threat to survival, with feral animals and alien plant species also of concern.

4.7 

Charpeniera tomentosa var. tomentosa

Slender to robust trees 6–12 m tall with densely brown tomentose new growth and stiff, coriaceous obovate to elliptic leaves. The marginal veins of the leaves are noticeably thickened. Occurs in mesic forest at elevations of 110–770 m, primarily in the Wai‘anae Mountains on O‘ahu but also known from Moloka‘i, Lāna‘i, Maui, and Hawai‘i. Two varieties are recognized, with var. tomentosa from the Wai‘anae Mountains pictured. The var. maakuaensis is known only from windward gulches of the Ko‘olau Mountains on O‘ahu.

Chenopodium—A large genus of about 100 species of often weedy herbs or shrubs, with 1 morphologically variable species endemic in the Hawaiian Islands. The name is from the Greek chen, goose, and pous, foot, because many species have leaves with this shape. At least 3 of 5 introduced species have become widely naturalized in the Islands.

4.9 

Chenopodium oahuense—high-elevation form

Scentless to weakly scented shrubs that are sometimes arborescent in growth, stems erect or prostrate. Leaves thick and somewhat fleshy, mealy pubescent, broadly deltate to rhombic, 1–3 cm in length, usually 3-lobed. Flowers in small dense clusters grouped into a larger terminal panicle. Occurs in a range of dry habitats over an elevational range from sea level to 2,520 m. Habitats include coastal, dry forest, and subalpine shrubland. Occurs on Lisianski, Laysan, French Frigate Shoals, Necker, Nīhoa, and all of the main islands. This species is highly variable in habit, leaf size, and leaf margin.

Nototrichium—An endemic genus of trees and shrubs represented by 3 species, 1 common and 2 at risk. Leaves opposite, with entire margins and dense stiff hairs on the leaf surfaces. The name is derived from the Latin, nota, remarkable, and the Greek, tricho, hair, in reference to the dense hairiness of the inflorescences.

Nototrichium sandwicense (A. Gray) Hillebr. kulu‘ī

ENDEMIC | SECURE

4.11 

Nototrichium sandwicense

Shrubs or small trees 1–4 m tall, usually branched from the base. Leaves densely pubescent, elliptic to ovate, 3–10 cm in length. Flowers in a stout, densely pubescent spike, 0.5–3.0 cm in length. Scattered

51

52

FLOWERING PLANTS

to occasionally common in dry open forest on exposed ridges and lava fields at elevations of sea level to 750 m on all of the main islands.

Anacardiaceae (SUMAC FAMILY)

Trees, shrubs, and woody vines. Some key features are the pinnate (or trifoliolate) leaves, the presence of vertical resin ducts in the bark, and drupaceous fruit. There are 73 genera and about 850 species distributed in the tropics, subtropics, Mediterranean climate regions, and temperate North America. Economically important food plants include mangoes, cashews, and pistachios. Many species of Rhus are planted as ornamentals. Oils, lacquers, and varnishes are obtained from other genera. Poison ivy and poison oak (Toxicodendron), well known members of this family, can cause severe dermatitis in sensitive individuals. In the Hawaiian Islands represented by the genus Rhus.

Rhus—A genus of about 200 species of

shrubs or small trees (or rarely lianas) in temperate and subtropical regions, especially South Africa. The name is derived from the Greek, rhous, which means sumac. Many species in this genus are economically important as sources of lacquers, tannins, waxes, and dyes. In the Hawaiian Islands there is 1 endemic species.

Rhus sandwicensis A. Gray neleau, neneleau, sumac ENDEMIC | SECURE

on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i. Most commonly seen along the road north of Hilo. The lightweight, coarse-textured wood is tough and was formerly used for saddle trees, yokes, and plows. A fungal disease has decimated many populations of this plant.

Apiaceae

(CARROT FAMILY)

Perennial, biennial, or annual herbs, shrubs, or rarely small trees. Some key features are aromatic herbage, the leaves usually with sheathing petioles, the inflorescence almost always in an umbel, and a distinctive fruit called a schizocarp. The family is cosmopolitan, with 428 genera and about 3,500 species worldwide. It is especially well represented in north temperate regions and tropical mountains. Economically important as food plants (e.g., celery, carrots, and parsnips) and as condiments (e.g., dill, anise, parsley, and fennel). Also some genera are cultivated as ornamentals. In the Hawaiian Islands represented by Peucedanum, Sanicula, and Spermolepis. All other genera are introduced, with the possible exception of Daucus pusillus, which is questionably indigenous.

Peucedanum—A genus of 100–120 species of caulescent perennial herbs with pronounced taproots; found in Eurasia, Africa, and the Hawaiian Islands. The name is from the Greek word for parsnip, peukedanum. Several species have medicinal properties as anti-inflammatory agents due to the presence of coumarin. In the Hawaiian Islands there is 1 endemic species.

Peucedanum sandwicense Hillebr. makou

ENDEMIC | VULNERABLE

4.12 

Rhus sandwicensis

Usually small trees 3–8 m tall with milky sap. Leaves odd-pinnately compound with 3–5 pairs of leaflets, terminal leaflet 9–14 cm in length. Flowers appear in dense, reddish-brown tomentose inflorescences in the fall. Grows in wet or dry disturbed places, generally at elevations of 150–750 m but occasionally higher,

4.13 

Peucedanum sandwicense

Stout, coarse, erect to sprawling perennial herbs, 50–100 cm tall. Plants with

a distinct odor of parsley. Leaves pinnately to ternate-pinnately compound, ovate to triangular-ovate, 10–25 cm in length. Flowers in terminal umbels. On windward cliffs at elevations of sea level to 250 m on Kaua‘i, O‘ahu (Wai‘anae Mountains), Moloka‘i, and Maui. There are 2 small populations in the Wai‘anae Range of O‘ahu, 3 on Moloka‘i, 10 on Kaua‘i, and 1 on Keopuka Rock, an islet close to Maui. These populations total 5,000–10,000 individuals, but only 2 populations with 61 individuals on O‘ahu. The root was used medicinally by Native Hawaiians

Sanicula—A genus of 39 species of perennial or biennial herbs arising from tuberous or fusiform roots, distributed worldwide in temperate or warm temperate regions, except Australia and New Zealand. The name is derived from the Latin sanare, which means to heal, as many of these plants have medicinal properties. The fruit of this genus is covered with tubercles or hooked bristles. In the Hawaiian Islands there are 4 endemic species.

Sanicula purpurea H. St. John & Hosaka snakeroot

ENDEMIC | ENDANGERED

4.14 

Sanicula purpurea

Somewhat stout, erect to ascending perennial herbs to 35 cm tall, arising from a massive many-branched caudex, stems many, trailing. Basal leaves numerous, kidney-shaped to orbicular, with 3–7 lobes, 2–8 cm in width. Flowers purplish. Scattered and rare on steep slopes and in open bogs at elevations of 700–1,460 m in the Ko‘olau Mountains, O‘ahu, and on West Maui. There are 5 known populations on O‘ahu with 24 individuals and 3 on Maui, with a combined total of about 210 individuals in 1996. Threats come from feral pigs and alien plant species, particularly the grass Axonopus fissifolius.

FLOWERING PLANTS

Sanicula sandwicensis A. Gray ENDEMIC | RARE

West Maui. Formerly known from open areas on Moloka‘i. In 1999 there were 12 known populations for a total of 2,000– 6,000 individuals.

Apocynaceae

(DOGBANE FAMILY)

4.15 

Sanicula sandwicensis

Erect perennial herbs up to 35 cm tall arising from a vertical fusiform root. Basal leaves numerous, palmately 3-parted into deeply lobed segments, 3–12 cm in width. Flowers yellow. Scattered in subalpine shrub and woodland at elevations of 2,000–2,600 m on the slopes of Haleakalā, East Maui, ‘Iao Valley on West Maui, and on Mauna Loa, Mauna Kea, and Hualālai, Hawai‘i.

Spermolepis—A genus of 5 species of annual herbs with slender taproots occurring in North America, Argentina, and the Hawaiian Islands. The name comes from the Greek sperma, seed, and lepis, scale, because of the spiny nature of the fruit. There is 1 endemic species in the Hawaiian Islands.

Spermolepis hawaiiensis H. Wolff ENDEMIC | ENDANGERED

Trees, shrubs, herbs, and lianas. Key features are the presence of milky sap, leaves mostly opposite or whorled, flowers that are contorted in bud, and seeds that often have a tuft of hairs. This family now includes the formerly separate Asclepiadaceae. In this new concept, it includes about 380 genera and 4,700 species; most are tropical, but the family is also well represented in temperate areas. Economically important for drug production (e.g., Nerium, Rauvolfia, and Strophanthus). Some taxa are a commercial source of rubber (e.g., Funtumia) and many are important ornamentals (e.g., Plumeria, Allamanda, Nerium, and Catharanthus). In the Hawaiian Islands represented by the endemic genus Pteralyxia and the indigenous genera Alyxia, Ochrosia, and Rauvolfia. Ornamentals such as the Madagascar periwinkle (Catharanthus roseus) and the yellow oleander (Thevetia peruviana) have become naturalized. Frangipani (Plumeria species) is extensively planted in the Hawaiian Islands.

Alyxia—A genus of 106 species ranging

in distribution from Malesia (a phytogeographical floristic region) to the West Pacific. The name is from the Greek alyktos, which means to be shunned or avoided. Members of this genus may be lianas or scandent to erect shrubs, some of which have medicinal value. In the Hawaiian Islands there is 1 indigenous species.

Alyxia stellata (J. R. Forst. & G. Forst.) Roem. & Schult. maile

4.18 

Alyxia stellata

Variable in habit from twining lianas to scandent or small erect shrubs. Leaves in threes, upper surface glossy, lanceolate to ovate or even suborbicular, 1–6 cm in length, fragrant when crushed. Drupes ovoid, often moniliform with 2–3 joints, black at maturity. Occasional to common in most vegetation types at elevations from 50–2,000 m. Once present on all of the main islands but no longer occurring on Ni‘ihau and Kaho‘olawe. The chemical coumarin accounts for the pleasant fragrance of the foliage, which is still used on festive occasions for decorations and lei throughout the Pacific Islands. Crushed maile was formerly used medicinally to cleanse open wounds. The Hawaiian plants represent an unnamed endemic variety. Listed in the Manual as Alyxia oliviformis.

Ochrosia—A genus of about 40 species of small trees native from the Mascarene Islands to Australia and the Pacific. The name is from the Greek ochros, which means pale yellow, referring to the yellow color of the fruit. Ochrosia elliptica is occasionally cultivated. In the Hawaiian Islands there are 4 endemic species.

Ochrosia haleakalae H. St. John hōlei

ENDEMIC | RARE

INDIGENOUS | SECURE

4.16 

Spermolepis hawaiiensis

Slender annual herbs to 20 cm tall, branches few. Leaves ternately compound with the ultimate divisions linear, 1–4 cm in length, basal leaves withering with age. Currently known only from small populations on Kaua‘i, O‘ahu, Lāna‘i, and

4.17 

Alyxia stellata—habit

4.19 

Ochrosia haleakalae

53

54

FLOWERING PLANTS

4.20 

Ochrosia haleakalae—fruit

Trees from 2–8 m tall. Leaves oblanceolate to elliptic, 3–4 per node, blades 6–15 cm in length. Flowers in relatively open inflorescences. Drupes yellow or purple at maturity. Scattered in dry to mesic forest at elevations of 700–1,200 m on East Maui and Hawai‘i (Pololū Valley, Kalōpā Gulch).

Trees from 3 to 8 m tall. Leaves oblanceolate to obovate, blades 11–22 cm in length, glossy dark green on upper surface. Fruit with small central and lateral wings at maturity. Rare on slopes and ridges in diverse mesic to sometimes wet forest at elevations of 250–610 m on Kaua‘i (Ku‘ia, Kalalau, Mahanaloa valleys, Hi‘i Mountains, Hā‘upu, Pāpa‘a, Wai‘oli-Waipā Ridge, Limahuli). There are less than 1,000 extant individuals, with habitat loss as a major threat.

ENDEMIC | VULNERABLE

Rauvolfia sandwicensis

Aquifoliaceae (HOLLY FAMILY)

kaulu

ENDEMIC | ENDANGERED

4.23 

Pteralyxia macrocarpa

Trees from 8 to 15 m tall. Leaves elliptic to obovate, blades 9–18 cm in length, glossy dark green on upper surface. Fruit with prominent lateral and central wings. Scattered and uncommon in species-rich mesic forest at elevations of 215–730 m on O‘ahu.

Rauvolfia—A pantropical genus of about

Pteralyxia kauaiensis

4.24 

From shrubs 1 m tall to trees 6 m or more. Leaves elliptic, in whorls of 3–5 per node, 6–16 cm in length. The bilobed fruit turn from green to purplish-black at maturity. Primarily found on ridges, slopes, and in gulches in mesic forest but also in remnant dry forest or on lava flows at elevations of 100–500 m or higher on all of the main islands except Kaho‘olawe. A variable plant throughout its range.

Pteralyxia kauaiensis Caum

4.22 

ENDEMIC | SECURE

kaulu

cies of trees from the Hawaiian Islands. The name is from the Greek pteron, wing, and the genus Alyxia. Unlike Alyxia, the inner wall of the bright red fruit is winged in this genus. Its affinities outside of the Hawaiian Islands are not known.

Pteralyxia kauaiensis—habit

hao

Pteralyxia macrocarpa (Hillebr.) K. Schum.

Pteralyxia—An endemic genus of 2 spe-

4.21 

Rauvolfia sandwicensis A. DC.

80 species of trees and shrubs. The name honors Leonhard Rauwolf, a sixteenth-century German who traveled in Asia. The alkaloid reserpine is derived from 2 species of this genus. The drug is used to treat high blood pressure and some forms of mental illness because of its tranquilizing effect. In the Hawaiian Islands there is 1 morphologically variable endemic species.

Trees and shrubs that are usually evergreen. Key features are leaves that are almost always spiral, the absence of stipules, usually dioecious with drupaceous fruit. There are 4 genera and 420 species that are cosmopolitan in distribution. Economically important for the hardwood of many species of Ilex. The same genus has many species that are popular ornamentals, including the Christmas holly. In the Hawaiian Islands represented by the indigenous genus Ilex.

Ilex—A genus of about 400 species of trees and shrubs that are cosmopolitan in distribution, especially in temperate regions. The name is from the Latin for holm oak, Quercus ilex. Some species are the source of teas (e.g., mate from Ilex paraguayensis), some are ornamentals (Ilex aquifolium and its cultivars, I. crenata and I. glabra), whose evergreen leaves and red berries are traditional Christmas decorations. In the Hawaiian Islands represented by 1 morphologically variable indigenous species. Two other species, Ilex aquifolium and I. paraguayensis, are sparingly naturalized in the Islands.

FLOWERING PLANTS

Ilex anomala Hook. & Arn. kāwa‘u, holly

INDIGENOUS | SECURE

Cheirodendron—A Polynesian genus of 6

species of trees and shrubs. The name is from the Greek cheiros, hand, and dendron, tree, because of the palmately compound leaves, often with 5 leaflets. In the Hawaiian Islands there are 5 endemic species, found primarily in wet forest, with 1 additional species in the Marquesas Islands. The plants have a strong carrot smell when cut or bruised.

Cheirodendron fauriei Hochr. ‘olapa, lapalapa

Trees from 2 to 10 m tall, leaves with 3 leaflets, broadly ovate, 3.5–7.0 cm in length, as wide or wider than long, veins of upper surface raised in dry leaves. Occurs in mesic and wet forest at elevations of 700– 1,300 m on Kaua‘i and O‘ahu. Two subspecies are recognized, differing in the shape of the leaflets. Pictured is subsp. platyphyllum found on O‘ahu, while the subsp. kauaiensis is restricted to Kaua‘i. The common name lapalapa is an onomatopoeia for the sound of the leaves when they flutter in the wind.

ENDEMIC | SECURE

Cheirodendron trigynum (Gaudich.) A. Heller olapa, lapalapa

ENDEMIC | SECURE

4.25 

Ilex anomala

Dioecious trees or occasionally shrubs from 5 to 12 m tall. Leaves variable in shape from oblanceolate to ovate, thick, coriaceous, 4–12 cm in length, with subentire margins. Common in mesic and wet forest and occasionally in bogs, typically at elevations of 600–1,400 m, on all of the main islands except Ni‘ihau and Kaho‘olawe. The wood was formerly used for saddle trees, canoe trimmings, and as anvils for kapa beating.

Araliaceae

(GINSENG FAMILY)

Trees, shrubs, lianas, woody epiphytes, and rarely herbs found widely in the neoand paleotropics and in some temperate regions. Key features are the usually umbellate inflorescence, commonly pinnately or palmately compound leaves, and 5-merous flowers. There are 39 genera and about 1,425 species. The family has limited economic importance. An exception is the ginseng root (Panax quinquefolius), which is widely sought worldwide for medicinal use, especially by the Chinese. English ivy (Hedera helix) is commonly cultivated as a ground cover in temperate regions, and species of Aralia, Polyscias, and Schefflera are widely planted as ornamentals in tropical and subtropical regions. In the Hawaiian Islands the family is represented by the indigenous genera Cheirodendron and Polyscias. Formerly the Hawaiian species of Polyscias were separated into the indigenous genus Reynoldsia and the endemic genera Munroidendron and Tetraplasandra, representing a single founding lineage.

4.26 

Cheirodendron fauriei

Trees 2–7 m tall, occasionally shrubby in growth. Leaves with 3 leaflets, coriaceous, 3–8 cm in length, about as wide as long, with their base obtuse to almost truncate. Occurs in mesic to wet forest at 650–1,250 m elevation on Kaua‘i (Kōke‘e, Kamo‘oloa Stream, Makaleha Mountains, Powerline Trail).

Cheirodendron platyphyllum (Hook. & Arn.) Seem. ‘olapa, lapalapa ENDEMIC | SECURE

4.27 

Cheirodendron platyphyllum subsp. platyphyllum

4.28 

Cheirodendron trigynum subsp. trigynum

Trees from 5 to 15 m tall. Leaflets vary from 3–5 per leaf, ovate to elliptic in shape, the largest leaflet 5–15 cm in length, usually twice as long as wide. This is by far the most variable species in morphology. Occurs in mesic and wet forest at elevations of 440–1,250 m. There are 2 subspecies, with the subsp. helleri restricted to Kaua‘i. The subsp. trigynum pictured is on all of the main islands except Kaua‘i and Kaho‘olawe. A bluish dye for kapa is made from the fruits, leaves, and bark of this plant. Also used in lei making.

Polycias—A genus of 159 species of paleotropical trees and shrubs with pinnately compound leaves and a gynoecium with 2 to many carpels. The Hawaiian species were once placed into the endemic genera Munroidendron and Tetraplasandra and the indigenous genus Reynoldsia. These and several other genera of Araliaceae have now been merged into a broader concept of Polyscias. The 11 Hawaiian species are all endemic and represent a single colonizing lineage.

55

56

FLOWERING PLANTS

Polycias hawaiensis (A. Gray) Lowry & G. M. Plunkett ‘ohe ENDEMIC | SECURE

Wai‘anae Mountains), Lāna‘i, Maui, and Hawai‘i. Listed in the Manual as Tetra­ plasandra kavaiensis.

Polyscias oahuensis (A. Gray) Lowry and G. M. Plunkett ‘ohe mauka

ENDEMIC | SECURE

4.29 

Trees up to 7 m tall with spreading branches and smooth gray bark. Leaves pinnately compound, 15–30 cm in length, leaflets 4–10. Long, pendulous inflorescences of pale yellow flowers make this species easy to recognize. Very rare and occurs only on Kaua‘i (Nounou Mountain, Nāpili Coast, Ha‘upu Ridge, Koai‘e Canyon) in mesic forest at elevations of 120–400 m. There are 15 extant populations, totaling about 200 trees. Listed in the Manual as Munroidendron racemosum.

Polyscias sandwicensis (A. Gray) Lowry & G. M. Plunkett ‘ohe, ‘ohe makai

Polycias hawaiensis

ENDEMIC | RARE

Trees 7–20 m tall, with diameters up to 30–60 cm. Leaves pinnately compound, 20–60 cm in length, leaflets 5–9. Fruit globose, ribbed when dry. Occurs in mesic to wet forest at 150–800 m elevation on Moloka‘i, Lāna‘i, Maui, and Hawai‘i. Listed in the Manual as Tetraplasandra hawaiensis.

Polyscias kavaiensis (H. Mann) Lowry & G. M. Plunkett ‘ohe ‘ohe ENDEMIC | SECURE

4.31 

Polycias oahuensis

Trees 3–10 m tall, highly variable. Leaves pinnately compound, 15–35 cm in length, glabrous on their lower leaf surface. Leaflets 7–15. Inflorescence umbellate, 2–3 times compound, fruit globose to ovoid, strongly ribbed when dry. Numerous forms of this plant have been described, resulting in a confusing taxonomy. Occurs in mesic to wet forest at elevations of 240–1,520 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in the Manual as Tetraplasandra oahuensis.

4.33 

Polycias sandwicensis—habit

4.34 

Polycias sandwicensis

Polyscias racemosa (C. N. Forbes) Lowry & G. M. Plunkett ENDEMIC | ENDANGERED

4.30 

Polycias kavaiensis

Trees 8–25 m tall. Leaves pinnately compound, 25–90 cm in length, lower leaf surface with stellate pubescence, leaflets 11–21. Inflorescence racemose, fruit globose to ovoid, 2–5 sided when dry. Occurs in mesic to wet forest at elevations of 650–1,600 m on Kaua‘i, where it is common, but rare on O‘ahu (northern

4.32 

Polycias racemosa

Trees up to 20 m tall and diameters up to 50–60 cm, with a spreading crown and thickish branch ends; wood with a yellowish resin. Leaves pinnately compound, deciduous in the dry season, 20–35 cm in length; leaflets typically 3. Fruit a globose drupe. Occurs in dry or occasionally mesic forest at elevations of 30–800 m on Ni‘ihau, O‘ahu, Moloka‘i, Lāna‘i, Maui, and Hawai‘i. Listed in the Manual as Reynoldsia sandwicensis.

FLOWERING PLANTS

Polyscias waialealae (Rock) Lowry & G. M. Plunkett ‘ohe kiko‘olā

ENDEMIC | SECURE

4.35 

Polycias waialealae

Trees 4–10 m tall, rarely to 12 m. Leaves pinnately compound, 20–35 cm in length, leaflets 7–13, thick and coriaceous. Inflorescence umbellate, 2–3 times compound. Fruit ovoid to cylindrical. Closely related to T. waimeae. Occurs in mesic to wet forest at 550–1,600 m elevation on Kaua‘i. Listed in the Manual as Tetraplasandra waialealae.

Polyscias waimeae (Wawra) Lowry & G. M. Plunkett ‘ohe kiko‘olā

ENDEMIC | SECURE

4.36 

Polycias waimeae

Trees 6–10 m tall. Leaves pinnately compound, 20–60 cm in length; leaflets 5–11, thick and coriaceous. Umbellate inflorescence once compound. Fruit large and globose. Closely related to T. waialealae. Occurs in wet forest at elevations of 550– 1,200 m on Kaua‘i. Listed in the Manual as Tetraplasandra waimeae.

Asteraceae

(SUNFLOWER FAMILY)

A cosmopolitan family of herbs, shrubs, climbers, and trees. The key feature of

the family is a head—that is, an inflorescence made up of multiple flowers borne on an enlarged base called the receptacle and surrounded by 1 or more series of subtending bracts called phyllaries. The inflorescence is typically composed of 2 types of flowers: ray and disk, although one or the other type may be absent; flowers set in a head subtended by phyllaries. The fruit is an achene usually crowned or appendaged with a pappus. The composites have been most recently organized into 42 tribes. Presently about 1,590 genera and 23,600 species are recognized, making this arguably the largest family of flowering plants. Although the family is cosmopolitan in distribution, the greatest diversity is in tropical and montane subtropical areas, with South Africa and the tropical Andes as centers of diversification. Economically the family is relatively unimportant considering its large size. Some are important food sources, such as Lactuca (lettuce), Cynara (artichoke), and Carthamnus (safflower oil); many are popular ornamentals, such as Chrysanthemum, Aster, Gazania, Calendula, Achillea, Helianthus, and Rudbeckia; some species yield significant insecticides, as pyrethrum from Tanacetum coccineum; some, such as Parthenium argentatum, yield rubber; others are obnoxious weeds, such as Bidens (beggar-ticks), Taraxacum (dandelion), Ambrosia (ragweed), and Sonchus (sow thistle). In the Hawaiian Islands, there are 13 native genera with 98 species. Endemic genera include Argyroxiphium, Dubautia, Hesperomannia, Lipochaeta, Remya, and Wilkesia. Indigenous genera include Adenostemma, Artemisia, Bidens, Keyserria, Melanthera, Pseudognaphalium, and Tetramolopium. There are at least 64 naturalized genera with about 90 naturalized species. Many introduced genera, such as Erechtites (fireweed), Hypochoeris (cat’s ear), Pluchea (Indian fleabane), and Sphagneticola (formerly Wedelia), have become serious pests.

Adenostemma—A genus of 24 species in tropical America, Africa, and Asia. All are annual or perennial herbs that are typically glandular pubescent. Heads discoid with the involucral bracts in 2 series, florets 10–60 per head, white or yellow. The name is from the Greek aden, meaning gland, and stemma, crown, referring to a crown of sticky pappus awns. These

sticky awns aid the plants in seed dispersal. Represented in the Hawaiian Islands by 1 indigenous species.

Adenostemma viscosum J. R. Forst. & G. Forst. kāmanamana

INDIGENOUS | SECURE

4.37 

Adenostemma viscosum

Perennial herbs 50–100 cm tall with ovate to rhombic leaves. Widespread from China, Malesia, Australia, Africa, Polynesia, and Micronesia. Uncommon in the Hawaiian Islands, where it occurs in mesic valleys and wet forests in shaded spots at elevations of 60–550 m on all of the main islands except Ni‘ihau and Kaho‘olawe. The Hawaiians used an infusion of the leaves of this plant to treat fevers.

Argyroxiphium—An endemic genus of 5

species native to bogs and alpine regions on Maui and Hawai‘i. Closely allied to Dubautia and Wilkesia, with which intergeneric hybrids occur both naturally and manmade. The name is from the Greek argyros, silver, and xiphos, sword, because of the shape and color of the leaves of the type species. Heads radiate to rarely discoid, ray florets 0–45 per head, disk florets 30–600 per head, heads extremely viscid-glandular and aromatic. Some species die after flowering, while others can reproduce by vegetative offshoots. One species, A. virescens, once occurred in mesic to wet forests on East Maui but is now considered extinct.

57

58

FLOWERING PLANTS

Argyroxiphium caliginis C. N. Forbes

Argyroxiphium kauense (Rock & M. Neal) O. Deg. & I. Deg.

‘Eke silversword

Ka‘ū silversword

ENDEMIC | RARE

ENDEMIC | ENDANGERED

4.38 

Argyroxiphium caliginis—habit

4.43 

4.41 

4.39 

Argyroxiphium grayanum—West Maui

Argyroxiphium caliginis

Argyroxiphium kauense

Erect, single-stemmed, or sparingly branched rosette shrubs arising from a woody stem up to 70 cm tall. The narrow sword-shaped leaves are usually silvery-silky but not as dense as in the Mauna Kea and Haleakalā species. Flower heads white to yellow. A rare plant in localized colonies in bogs or wet forest on Mauna Loa at elevations of 1,625–1,900 m. One collection is known from Hualālai. Populations have seriously declined in the last century to an estimated 400 individuals. Mouflon sheep have seriously impacted populations in the past.

Argyroxiphium sandwicense DC. ‘ahinahina, hinahina, silversword

Dwarf shrubs with a spreading, manybranched caudex. The silvery leaves are in a basal rosette. The corollas in this species are purplish. Occurs in the summit bogs of ‘Eke and Pu‘ukukui at elevations of 1,350–1,650 m on West Maui.

ENDEMIC | VULNERABLE/ENDANGERED

Argyroxiphium grayanum (Hillebr.) O. Deg. greensword

ENDEMIC | SECURE

4.42 

4.40 

Argyroxiphium grayanum—habit

Argyroxiphium grayanum—East Maui

Erect, branched or unbranched, or decumbent shrubs to 2 m. The swordshaped leaves are green and the corollas yellowish. Found in and around the summit bogs of West Maui and in wet forest above the Hāna Forest Reserve on East Maui at elevations of 1,200–2,050 m. West Maui plants tend to be smaller than those on East Maui.

Argyroxiphium sandwicense subsp. macrocephalum—habit 4.44 

FLOWERING PLANTS

recovering from a population low of only 40 individuals.

Artemisia mauiensis (A. Gray) Skottsb. ‘āhinahina, hinahina ENDEMIC | SECURE

Artemisia—A genus of about 400 species

of aromatic herbs and shrubs from the north temperate areas, western South America, and 1 species in South Africa. The finely dissected gray-green leaves of Hawaiian species are highly distinctive and easily recognized. Heads discoid, with involucral bracts in 2–3 overlapping ranks. The name honors the Greek goddess of the hunt, Artemis (also known as Diana). Economically important for the production of vermifuges, stimulants, and culinary herbs. The spirit absinthe is distilled from leaves of A. absinthium. In the Hawaiian Islands there are 3 endemic species representing a single colonizing lineage from Southeast Asia.

Argyroxiphium sandwicense subsp. macrocephalum 4.45 

Artemisia australis Less. ‘āhinahina, hinahina ENDEMIC | SECURE

4.48 

Artemisia mauiensis

Argyroxiphium sandwicense subsp. sandwicense

Faintly aromatic subshrubs branched at the base with numerous erect to decumbent stems up to 40 cm in length. Leaves broadly ovate in outline, 3–5 cm in length, 2–3 pinnatifid into filiform segments less than 1 mm in width, densely white-tomentose on both surfaces. Known from exposed faces of volcanic rock in subalpine shrubland at elevations of 1,900–2,300 m elevation on Halea­ kalā, East Maui. Occasionally this species appears to hybridize with A. australis, where the two occur together.

Single-stemmed (rarely branched) shrubs with a dense basal rosette of silvery leaves. Flower heads pink to wine red, changing to yellow at their base. The large flowering stalk, usually borne during the summer months, may be up to 2 m tall. The plants die after flowering. Confined to the subalpine and alpine regions of Haleakalā on East Maui and Mauna Kea on Hawai‘i at elevations of 2,125–3,750 m. The var. macrocephalum from Maui has larger flowers than the form on Hawai‘i. Once common, these magnificent plants were on the verge of extinction earlier in the century due to the ravages of man, goats, and insects. Several introduced insects are having direct and indirect negative impacts on their survival. The var. sandwicense is rare and locally restricted on Mauna Kea but is

Bidens—A large genus of cosmopolitan herbs and shrubs with about 200 recognized species. The opposite leaves typically simple to pinnately compound, with serrate margins. Flower heads radiate or discoid with involucral bracts in 2 series, ray florets few and typically yellow or white, disc flowers yellow to orange. The name is from the Latin bi, two, and dens, teeth, because of the nature of the pappus awns. The obnoxious weeds Bidens pilosa and B. alba, beggar-ticks or Spanish needles, are naturalized in the Hawaiian Islands. Bidens alba has rapidly now replaced B. pilosa as the most common roadside weed. The retorsely barbed awns on the fruit of this species readily attach themselves to fur or feathers, making this an effective mechanism for bird and mammal dispersal. In the Hawaiian

4.46 

4.47 

Artemisia australis

Faintly to strongly aromatic subshrubs, diffusely branching at the base, with numerous erect to decumbent stems. Leaves broadly ovate in outline, 5–7 cm in length, finely dissected, leaf segments narrow to lanceolate but never filiform, gray-green, upper surface lightly pubescent, lower surface white tomentose. A variable species in the Hawaiian Islands, especially in the shape of the leaves. Occurs on exposed cliff faces on the windward sides of all of the main islands at elevations of sea level to 1,065 m.

59

60

FLOWERING PLANTS

Islands there are 19 endemic species, which have arisen from a single colonizing lineage. All are interfertile. Almost all of the endemic Hawaiian species show loss of a long-distance dispersal ability.

Bidens forbesii Sherff

Bidens menziesii (A. Gray) Sherff

ENDEMIC | SECURE

ENDEMIC | VULNERABLE/SECURE

ko‘oko‘olau, ko‘olau, beggar’s tick

ko‘oko‘olau, ko‘olau, beggar’s tick

Bidens amplectens Sherff

ko‘oko‘olau, ko‘olau, beggar’s tick ENDEMIC | RARE

4.49 

Bidens amplectens

4.51 

Bidens forbesii subsp. kahiliensis

Erect perennial or facultative annual herb, 0.5–1.3 m tall. Leaves pinnately compound, 9–15 cm in length, leaflets 3–5. Flowers yellow. Growing in low shrubland on windward cliffs at 50–220 m elevation in the Wai‘anae Mountains between Kawaihāpai and Ka‘ena Point, O‘ahu. Threats to survival come from alien plant species and fire.

Erect suffrutescent perennial herb, 0.7–2.0 m tall. Leaves pinnately compound or some simple, 11–36 cm in length, leaflets (1) 3–7. Flowers yellow. Infrequent to locally common on coastal bluffs and cliffs and mesic forests from sea level to 1,000 m elevation on Kaua‘i. The subsp. forbesii is widespread on Kaua‘i, while the subsp. kahiliensis is restricted in distribution to Kāhili and Kapalaoa peaks on Kaua‘i.

Bidens cosmoides (A. Gray) Sherff

Bidens hillebrandiana (Drake) O. Deg.

ENDEMIC | VULNERABLE

ENDEMIC | SECURE

ko‘oko‘olau, ko‘olau, po‘olā nui, beggar’s tick

ko‘oko‘olau, ko‘olau, beggar’s tick

4.53 

Bidens menziesii subsp. filiformis

Erect, sparingly branched shrubs 0.8–4.0 m tall. Leaves bipinnately divided into long linear segments, 4–26 cm in length. Flowers yellow. There are 2 recognized subspecies. Pictured is the subsp. filiformis with narrower leaves that is common on cinder slopes and subalpine forest at 750–2,200 m elevation on the leeward slopes of Mauna Loa and Mauna Kea and in the saddle area between the mountains. The subsp. menziesii occurs on arid leeward slopes and cliffs in shrubland at elevations of 200–750 m on Moloka‘i and West Maui. Leaves of this and other native species of Bidens were made into a tea by the Native Hawaiians that was considered a tonic or blood purifier. They were also used to treat thrush, a fungus disease formerly common in children.

Bidens molokaiensis (Hillebr.) Sherff 4.52 

4.50 

Bidens cosmoides

Scandent, suffrutescent perennial herbs 1.5–2.0 m tall. The unique flower heads, with large yellow ray flowers, are unlike any other species in the genus and may represent an adaptation to pollination by birds. Scattered and formerly common in diverse mesic forest at 750–1,200 m elevation on Kaua‘i. Threats to survival come from alien plant species and fire.

Bidens hillebrandiana subsp. polycephala

Erect or trailing suffrutescent perennial herbs about 20–30 cm tall. Leaves typically pinnately or bipinnately compound, slightly fleshy, 3.5–8.0 cm in length, leaflets 3–9. Flowers yellow. Occurs on coastal bluffs and sea cliffs at elevations of sea level to 100 m on Moloka‘i, East Maui, and Hawai‘i. There are 2 subspecies, with subsp. polycephala from the windward coasts of Moloka‘i and East Maui pictured. The subsp. hillebrandiana occurs on the Kohala Coast of Hawai‘i.

ko‘oko‘olau, ko‘olau, beggar’s tick ENDEMIC | VULNERABLE

4.54 

Bidens molokaiensis

FLOWERING PLANTS

Trailing perennial herb, 10–30 cm in height. Leaves simple or trifoliate, usually 3–7 cm in length. Flowers yellow. Scattered in occurrence on coastal cliffs from sea level to 250 m elevation on windward Moloka‘i. Once occurring on Diamond Head on O‘ahu but now extinct there. Threats to survival come from alien plant species, feral animals, and fire.

Bidens sandvicensis Less.

ko‘oko‘olau, ko‘olau, beggar’s tick ENDEMIC | RARE/SECURE

lateral branches. Leaves pinnately or bipinnately compound, 9–30 cm in length, young leaves with a serrated margin and with a white, golden, or rust-colored pubescence. Flowers yellow. The achenes of this species are strongly coiled or twisted, unlike those of the other Hawaiian Bidens. Common on dry ridges and in wet forest at elevations of 300–1,200 m on O‘ahu, primarily in the Wai‘anae Mountains but also in the northwestern Ko‘olau Mountains.

Dubautia arborea (A. Gray) D. D. Keck na‘ena‘e, kūpaoa ENDEMIC | RARE

Bidens valida Sherff

ko‘oko‘olau, ko‘olau, beggar’s tick ENDEMIC | SECURE

4.58  4.55 

Bidens sandvicensis subsp. confusa

Erect, suffrutescent perennials 0.5–1.0 m tall. Leaves pinnately compound or rarely bipinnately compound, 5–19 cm in length; leaflets 3–7. Flowers yellow. Common on ridges in disturbed shrubland and in diverse mesic forest at elevations of 150–900 m on Kaua‘i and O‘ahu. There are 2 subspecies, with subsp. sandwicensis from Kaua‘i and O‘ahu. Pictured is the rare subsp. confusa, with narrower leaves, which occurs only in Waimea Canyon, Kaua‘i.

Bidens torta Sherff

ko‘oko‘olau, ko‘olau, beggar’s tick ENDEMIC | SECURE

4.56 

Bidens torta

Erect suffrutescent perennial herbs, 1.0– 2.5 m tall with horizontal or ascending

4.57 

Bidens valida

Erect suffrutescent perennial herbs, 0.7–1.5 m tall with horizontal lateral branches. Leaves pinnately compound, 9–20 cm in length; leaflets typically 3–5. Scattered on exposed windswept ridges in mesic to wet forest at elevations of 350–900 m on the summits of Kāhili and Hā‘upu Ridge, Kaua‘i.

Dubautia arborea

Large shrubs to small trees up to 6 m tall, trunks up to 40 cm in diameter. Leaves in whorls of 3, chartaceous, foetid when fresh, usually with 5 main veins. Flowers yellowish orange. Once common but now only locally abundant in subalpine shrub­ land and arid alpine sites at elevations of 2,120–3,100 m on the upper slopes of Mauna Kea (Pu‘ulā‘au), Hawai‘i.

Dubautia carrii B. G. Baldwin & Friar na‘ena‘e, kūpaoa

ENDEMIC | VULNERABLE

Dubautia—An endemic genus of sub-

shrubs, shrubs, trees, and 1 liana, with 26 recognized species. Leaves opposite, alternate, or whorled. Heads discoid, flowers yellow or white. Five new species have been described since the publication of the Manual. Diversity and endemism are notable on Kaua‘i, where 11 of the 13 Dubautia species present are restricted to the island. Part of the silversword alliance with Argyroxiphium and Wilkesia. Natural hybrids occur, including a generic cross between the Haleakalā silversword and Dubautia menziesii where the two grow together. The name honors J. E. Dubaut, an officer in the French Royal Marines.

4.59 

Dubautia carrii

61

62

FLOWERING PLANTS

Compact to diffusely branched shrubs, typically 1.5–3.0 m tall. Leaves opposite, broadly to narrowly elliptic. With 3–5 main veins, apex toothed. Flowers yellow to yellow-orange. Closely resembles D. linearis but differs in leaf form and habitat. Occurs in mesic to wet forest and shrubland at elevations of 450–1,150 m on eastern Moloka‘i (Pu‘u Ka‘eha).

subsp. ciliolata from Kīlauea, Mauna Loa, and Hualālai and subsp. glutinosa from Mauna Kea.

Dubautia herbstobatae G. D. Carr na‘ena‘e, kūpaoa

ENDEMIC | ENDANGERED

Dubautia ciliolata (DC.) D. D. Keck na‘ena‘e, kūpaoa

ENDEMIC | SECURE

Openly branched shrubs or small trees, 1.8–3.5 m tall, stems with stiff hairs. Leaves opposite, narrowly elliptic to oblanceolate, with 7 primary veins. Flowers greenish white. This taxa was previously treated within a broad concept of D. laxa subsp. hirsuta, but it differs in having white rather than yellow flowers and the presence of stiff hairs on the lower leaf surface. Rare in wet forests at elevations of 1,205–1,235 m on the southern rim of the Kalalau Valley, Kaua‘i. There is a single known population with 26 individuals.

Dubautia knudsenii Hillebr. na‘ena‘e, kūpaoa

ENDEMIC | RARE/SECURE

4.62 

4.60 

Dubautia ciliolata subsp. ciliolata

Dubautia herbstobatae

Small speading shrubs up to 50 cm tall. Leaves opposite or rarely ternate, coriaceous, upper surface lustrous, with 1 or rarely 3 main veins. Flowers yellow-orange. Rare on dry ridgetops at elevations of 580–915 m in the northwestern Wai‘anae Mountains, O‘ahu (‘Ōhikilolo, Kamaile‘unu Ridge). From a population low of less than 100 indivduals, restoration efforts and removal of goats from the area have been successful, resulting in more than 2,000 individuals today. Threats come from erosion, fire, and human activities.

Dubautia kalalauensis B. G. Baldwin & G. D. Carr na‘ena‘e, kūpaoa ENDEMIC | ENDANGERED

4.64 

Dubautia knudsenii subsp. knudsenii

Shrubs or small trees to 6 m tall. Leaves opposite, elliptic to elliptic-oblanceolate, faintly 5–9 nerved. An interesting species in which the flower heads are usually reflexed, flowers yellow. Occurs in mesic to wet forests and bogs at elevations of 550– 1,275 on Kaua‘i. Three subspecies are recognized, with subsp. knudsenii pictured.

Dubautia latifolia (A. Gray) D. D. Keck koholāpehu, kūpaoa, na‘ena‘e ENDEMIC | ENDANGERED

4.61 

Dubautia ciliolata subsp. glutinosa

Erect, often many-branched shrubs to 1.8 m tall. The smoothly varnished leaves are usually in whorls of 3 except for the plants occurring on Mauna Kea, which are opposite and sticky-glandular. Flowers yellow. Often hybridizes with D. scabra where the two grow together. Common in open, dry shrubland, dry ‘ō‘hia forest and subalpine forest at elevations of 900– 3,200 m on Hawai‘i. Pictured are both

4.63 

Dubautia kalalauensis

4.65 

Dubautia latifolia

FLOWERING PLANTS

Diffusely branched lianas with slender stems up to 8 m or more long. The widely spaced, elliptic leaves are finely reticulate, unlike any others in the genus, and petiolate. Flowers yellow. A rare plant occurring in diverse mesic forest at elevations of 975–1,200 m in the Kōke‘e area of Kaua‘i. The aggressive introduced vine Passiflora tarminiana and other alien plants threaten the survival of this species. Less than 200 extant individuals are known.

Many-branched diffuse shrubs up to 3 m tall. Leaves may be ternate, opposite, or alternate, with variable form sometimes occurring on a single branch, obscurely 3–5 veined. Flowers yellowish orange. Occurs from dry shrubland to wet forest at 450–2,500 m elevation on Moloka‘i, Lāna‘i, Maui, and Hawai‘i. There are 2 named subspecies, with subsp. linearis from Lāna‘i and Maui pictured. The subsp. hillebrandii is restricted to Hawai‘i.

Dubautia laxa Hook. & Arn.

Dubautia menziesii (A. Gray) D. D. Keck

ENDEMIC | SECURE

ENDEMIC | SECURE

na‘ena‘e pua melemele, na‘ena‘e, kūpaoa

na‘ena‘e, kūpaoa

Dubautia paleata A. Gray

kūpaoa, na‘ena‘e pua kea, na‘ena‘e ENDEMIC | SECURE

4.70 

Dubautia paleata

Shrubs to 2.5 m tall. The leaves opposite or in a whorl of 3, usually exhibing notable pubescence and variable in shape and length (3.5–20.0 cm), with 7–15 primary veins. Flowers white. A species found primarily in and around the margins of bogs at 1,100–1,550 m elevation on Kaua‘i.

Dubautia plantaginea Gaudich. 4.68  4.66 

Dubautia laxa subsp. hirsuta

Shrubs up to 5 m tall. Leaves opposite, elliptic to oblong-oblanceolate, usually 5–7 veined. Flowers yellow. A highly variable species with 4 distinct subspecies that differ in the number of flowers in the heads and shape and pubescence of the leaves. Occurs in wet forests and bogs at elevations of 360–1,700 m on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui. There are 4 named subspecies. Pictured is subsp. hirsuta, which occurs on Kaua‘i, O‘ahu, and Lana‘i.

Dubautia menziesii

na‘ena‘e, kūpaoa

ENDEMIC | SECURE/ENDANGERED

Small to large shrubs, to 2.5 m tall. Leaves in whorls of 3, opposite or alternate, usually imbricate, coriaceous, and mildly balsam scented when fresh, typically with 3 primary veins. Flowers yellowish orange. A common plant of subalpine shrubland and alpine desert at elevations of 1,800– 3,075 m on Haleakalā, East Maui.

Dubautia microcephala Skottsb. na‘ena‘e, kūpaoa ENDEMIC | RARE

4.71 

Dubautia plantaginea subsp. magnifolia

4.72 

Dubautia plantaginea subsp. plantaginea

Dubautia linearis (Gaudich.) D. D. Keck na‘ena‘e, kūpaoa

ENDEMIC | SECURE

4.69 

4.67 

Dubautia linearis subsp. linearis

Dubautia microcephala

Shrubs to small trees up to 4 m tall with relatively few spreading branches. Leaves opposite, linear-lanceolate to linear-oblanceolate, margins toothed along their upper half, with 11–17 primary veins. Flowers yellow, reflexed. Occurs in mesic forest at 825–1,275 m elevation on Kaua‘i.

63

64

FLOWERING PLANTS

Shrubs or small trees up to 7 m tall. Leaves opposite, 8–26 cm in length, with 7–15 primary veins. A variable plant with respect to the size and surface characteristics of the leaves. The large inflorescence is pyramidal in shape. Flowers yellow, turning purplish with age. Distributed in mesic to wet forest at elevations of 300–2,100 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Three subspecies are recognized that differ in pubescence, leaf size, and nature of the branching of the inflorescence. The subsp. magnifolia is rare on Kaua‘i and known from just 2 populations, while subsp. plantaginea occurs over the range of the species and is secure. The subsp. humilis is a rare dwarf taxa restricted to Black Gorge on West Maui.

Dubautia platyphylla (A. Gray). D. D. Keck na‘ena‘e, kūpaoa ENDEMIC | RARE

Dubautia raillardioides Hillebr.

Dubautia scabra (DC.) D. D. Keck

ENDEMIC | SECURE

ENDEMIC | SECURE

na‘ena‘e ‘ula, kūpaoa

na‘ena‘e, kūpaoa

4.76 

4.74 

Dubautia raillardioides

Somewhat scandent to partly trailing shrubs up to 3 m tall, with thick but fleshy trunks. Leaves are in whorls of 3, lanceolate to oblong-oblanceolate, 9–25 cm in length, glabrous. Flowers white. Closely related to Dubautia paleata. Occurs in wet forest, usually at the margins of bogs, at elevations of 600–1,375 m on Kaua‘i.

Dubautia reticulata (Sherff) D. D. Keck na‘ena‘e, kūpaoa

Dubautia scabra subsp. scabra

Suberect or decumbent, spreading or mat-forming shrubs with narrowly linear leaves. Often found on recent lava flows on Hawai‘i at elevations from 75 to 2,500 m. Flowers white. A conspicuous plant in the Kīlauea area and also along the Saddle Road between Mauna Kea and Mauna Loa. Pictured is subsp. scabra, which occurs on East Maui and Hawai‘i. The subsp. leiophylla is found in wet forest on Moloka‘i, Lāna‘i, Maui, and Hawai‘i (Kohala Mountains) and does not form mats.

Dubautia syndetica G. D. Carr & Lorence na‘ena‘e, kūpaoa ENDEMIC | RARE

ENDEMIC | RARE

4.73 

Dubautia platyphylla

Large shrubs up to 3.5 m tall, with thick stems. Leaves opposite or occasionally in whorls of 3, ovate-lanceolate in shape, with 7–11 primary veins. Young shoots have a citruslike aroma. Flowers yellowish orange. Occurs in ravines and on cinder slopes with subalpine shrubland at elevations of 1,900–2,750 m on East Maui.

4.75 

Dubautia reticulata

Large shrubs or small trees up to 8 m tall, with broad trunks. Leaves opposite and spreading, typically with 5–7 primary veins. Flowers yellowish orange. Occurs in wet forest at elevations of 1,575–2,300 m on East Maui. The current range is much reduced from where it once occurred, and it is most common today on the east side of Ko‘olau Gap. Hybridizes with D. waianapanapaenis.

4.77 

Dubautia syndetica

Shrub 1–2 (3) m tall. Leaves opposite, 4–16 cm in length. The diagnostic features of the species include uniseriate, somewhat coalescent receptacular bracts on the receptacle, glandular peduncles and corolla tubes, and achenes with matted pubescence. Restricted to the headwaters of Wahiawa Stream, Kaua‘i, in diverse lowland wet forest, at elevations of 680–950 m.

FLOWERING PLANTS

Dubautia waialealae Rock na‘ena‘e, kūpaoa

ENDEMIC | ENDANGERED

4.78 

Dubautia waialealae

Small shrubby trees or shrubs 2–3 m tall. The leaves are elliptic to elliptic-ovate. The flowers in this species are erect and in clusters of 4–5. A rare plant on slopes and ridges in mesic to wet forest at elevations of 360–750 m in the Wai‘anae Mountains of O‘ahu and West Maui. Populations have been declining and currently known from 5 locations with just 14 individuals on O‘ahu and a single plant growing in the ‘Īao Valley of West Maui. Threats come from habitat loss, feral pigs, and alien plant species.

Hesperomannia lydgatei C. N. Forbes ENDEMIC | ENDANGERED

Low compact shrub, cushionlike in growth form, 10–50 cm tall. Leaves in whorls of 3–4 per node, closely imbricate, 1–3 cm in length, 3–5 toothed at the apex. Flowers yellow. Limited in occurrence to the summit bog of Mount Wai‘ale‘ale at an elevation of 1,450–1,575 m. The population is estimated to number about 3,000 individuals.

flowers are in clusters of 4–5 but nodding at maturity. Rare and local in dense wet forest at elevations of 600–750 m along the Wahiawa and Wai‘oli streams, Kaua‘i. Known from 8 localities with a total of less than 300 individuals; threats are from invasive alien plant species.

Keysseria—A genus of perennial herbs or small shrubs with about 12 species distributed from Malesia to the Hawaiian Islands. The name honors Christian Keysser, who collected the type specimen in New Guinea. In the Hawaiian Islands represented by 3 endemic species, mostly occurring in high-elevation boggy or wet places. These species were formerly placed in the genus Lagenifera.

Keysseria maviensis (H. Mann) Cabrera hōwaiaulu

ENDEMIC | VULNERABLE

Gnaphalium—Hawaiian species formerly placed in the genus Gnaphalium are now included in Pseudognaphalium.

Hesperomannia—An endemic Hawaiian genus of shrubs or small trees, with 3 species known, all of them quite rare. Leaves simple, alternate. Heads large, discoid, flowers yellow to yellowish brown. The name honors a Western botanist, Horace Mann (hesperos, the Greek word for west). Although once placed in the tribe Mutisieae, molecular data show a relationship to African Vernonieae.

4.80 

Hesperomannia lydgatei—habit

Hesperomannia arbuscula Hillebr. ENDEMIC | ENDANGERED

4.82 

4.81 

4.79 

Hesperomannia arbuscula

Hesperomannia lydgatei

Small trees 2–3 m high with elliptic-obovate to broadly oblanceolate leaves. The

Keysserria maviensis

Perennial herbs or subshrubs that are usually branched at the base. The leaves are generally in a basal rosette. This is the only Hawaiian species with conspicuous white ray flowers. Occurs in bogs, sometimes as an epiphyte, and in wet forest at elevations of 915–1,980 m on Maui and known from a single collection on Moloka‘i (Kamapamoa Ridge). Listed in the Manual as Lagenifera maviensis.

Lagenifera—Hawaiian species formerly placed in this genus are now placed in Keysseria.

65

66

FLOWERING PLANTS

Lipochaeta connata (Gaudich.) DC.

Spreading to trailing suffrutescent perennial herbs, with branches 0.5–1.5 m in length. Leaves herbaceous to somewhat leathery in texture, lanceolate-linear in shape, 4–10 cm in length, margins serrate. Infrequent to locally common in dry coastal habitats and dry shrubland from sea level to 400 m elevation on Ni‘ihau, O‘ahu, and West Maui. There are 2 named subspecies, with the subsp. lobata on Ni‘ihau, O‘ahu, and West Maui.

ENDEMIC | SECURE

Lipochaeta rockii Sherff

Lipochaeta—An endemic Hawaiian genus

of 6 species of somewhat shrubby perennial herbs and 1 annual found in mostly dry or mesic localities. Stems rounded or weakly angled; disk flowers 4-merous. A group of 14 species considered as a subgenus of Lipochaeta in the Manual is now treated as the indigenous genus Melanthera.

nehe

nehe

ENDEMIC | SECURE

6–12 cm in length, fleshy. Scattered along the coasts at elevations from sea level to 100 m on all of the main islands except Lāna‘i. Rare on O‘ahu.

Melanthera—A genus of about 35 species of perennial herbs with a pantropical distribution. The Hawaiian species were considered to form a subgenus of Lipochaeta in the Manual. These species differ from Lipochaeta in having conspicuously angled stems, petiolate rather than sessile leaves, and disk flowers 5-merous rather than 4-merous. The name comes from the Greek melan, black, and the Latin anthera, anther. In the Hawaiian Islands there are 14 endemic species representing a single colonizing lineage.

Melanthera integrifolia (Nutt.) W. L. Wagner & H. Rob. nehe

ENDEMIC | SECURE

4.85  4.83 

Lipochaeta connata subsp. connata

Erect suffrutescent perennial herbs, 10–20 cm tall. Leaves opposite or rarely in whorls of 3, 8–19 cm or more in length, ovate to broadly elliptic or lanceolate. Scattered in remnant dry forests at 2–400 (1,160) m elevation on Ni‘ihau, Kaua‘i, and West Maui. There are 2 named subspecies, with the subsp. connata on Kaua‘i and West Maui.

Lipochaeta lobata (Gaudich.) DC. nehe

Lipochaeta rockii

Spreading to erect suffrutescent perennial herbs, with branches 0.5–1.0 m in length. Leaves ovate to deltate in shape, 4–10 cm in length, with upper and lower surfaces pubescent. Infrequent to locally common in coastal areas of dry forests, particularly in disturbed areas and along the margins of lava flows, at 15–550 m elevation on Moloka‘i, Kaho‘olawe, and Maui, with 1 old collection from Hawai‘i.

Lipochaeta succulenta (Hook. & Arn.) DC. nehe

ENDEMIC | SECURE

4.87 

Melanthera integrifolia

Prostrate suffrutescent perennial herbs, rooting along their stems. Leaves somewhat fleshy, oblong to spathulate in shape, 1–3 cm in length. Scattered to locally common along the coasts of all of the main islands and on Kure and Laysan atolls. Listed in the Manual as Lipochaeta integrifolia.

ENDEMIC | SECURE/ENDANGERED

Melanthera kamolensis (O. Deg. & Sherff) W. L. Wagner & H. Rob. nehe ENDEMIC | ENDANGERED

4.86 

4.84 

Lipochaeta lobata subsp. lobata

Lipochaeta succulenta

Trailing suffrutescent perennial herbs, rooting along the lower surface, occasionally erect. Leaves linear to elliptic-ovate,

4.88 

Melanthera kamolensis

FLOWERING PLANTS

Trailing to scandent suffrutescent perennial herbs with stems rooting along their lower surfaces, stems 0.3–3.0 m in length. Leaves linear to deltate in shape, 3.0–6.5 cm in length, both surfaces with slender, stiff hairs. A very rare plant known only from remnant dry forest at about 250 m elevation on southeastern Maui (Kamole and Kepuni gulches). The single pure population in Kamole Gulch consists of only 25 mature individuals, while the second population of about 100 mature individuals is hypothesized to be hydridized with Lipochaeta rockii. Threats come from feral grazing animals, alien plant species, and fire. Listed in the Manual at Lipochaeta kamolensis.

Trailing suffrutescent perennial herbs with stems rooting along their lower surface. Leaves deltate, 2–10 cm in length, entire to lobed and pinnatifid. Known from diverse mesic forest at elevations of 300–400 m on Kaua‘i. Two endangered varieties occur that differ in leaf shape and flower length. Pictured is subsp. micrantha, known from Olokele Canyon and Hanapēpē Valley. Listed in the Manual as Lipochaeta micrantha.

Melanthera remyi (A. Gray) W. L. Wagner & H. Rob. nehe

Suffrutescent perennial herbs with erect to ascending stems to 3 m tall. Leaves narrowly deltate, 3–10 cm in length, usually with 2 basal lobes. A polymorphic species. Occurs in dry forest and introduced grassland at elevations of 550–1,800 m in the North Kona and South Kohala districts of Hawai‘i. Listed in the Manual as Lipochaeta subcordata.

Melanthera venosa (Sherff) W. L. Wagner & H. Rob. nehe ENDEMIC | ENDANGERED

ENDEMIC | RARE

Melanthera lavarum (Gaudich.) W. L. Wagner & H. Rob. nehe

ENDEMIC | SECURE

4.93  4.89 

Melanthera lavarum

Erect suffrutescent perennial herbs, stems up to 2 m in length. Leaves gray to grayish green, linear to elliptic or narrowly ovate, 3.0–6.5 cm in length. An exceedingly variable plant in leaf size and form. Scattered to occasionally common in coastal habitats or in dry forest from sea level to 550 m elevation on Moloka‘i, Lāna‘i, Maui, Kaho‘olawe, and northwestern Hawai‘i. Listed in the Manual as Lipochaeta lavarum.

Melanthera micrantha (Nutt.) W. L. Wagner & H. Rob. nehe

4.91 

Melanthera remyi

Annual herbs with erect stems to 60 cm tall. Leaves thin and ovate in shape, 3–7 cm in length. The only annual species in the genus. Local in moist sites in dry forest at elevations of 30–180 m in the northwestern Wai‘anae Mountains, O‘ahu. Listed in the Manual as Lipochaeta remyi.

Melanthera subcordata (A. Gray) W. L. Wagner & H. Rob. nehe

ENDEMIC | SECURE

Melanthera micrantha subsp. micrantha

Suffrutescent perennial herbs with arching stems. The dimorphic leaves are sometimes pinnately dissected, while others are coarsely serrate, 2–3 cm in length, both surfaces with slender, stiff hairs. Closely related to Melanthera subcordata. Rare in dry shrubland on cinder cones at elevations of 730–915 m on northwestern Hawai‘i (Nohonaohae, Holoholokū, Heihei cinder cones, Pu‘upā, South Kohala District). There are 6 populations with a total of about 3,000 individuals. Threats come from feral grazing animals, alien plant species, and fire. Listed in the Manual as Lipochaeta venosa.

Pseudognaphalium—A genus of about 100

ENDEMIC | ENDANGERED

4.90 

Melanthera venosa

4.92 

Melanthera subcordata

species of annual or perennial herbs with a distribution centered in North America. They are commonly called cudweeds, and many are aromatic. The name comes from the Greek pseudo, appearing similar to, and gnaphalon, lock of wool, since many of the species are woolly pubescent. Some species are cosmopolitan weeds. One endemic species with 4 distinct varieties is known from the Hawaiian Islands, as well as several naturalized species.

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FLOWERING PLANTS

Pseudognaphalium sandwicensium (Gaudich.) A. Anderb. ‘ena‘ena, cudweed, everlasting

ENDEMIC | SECURE/VULNERABLE

Weakly erect to scandent shrubs 1–4 m tall. Leaves lanceolate to elliptic-lanceolate or even broadly ovate, 4.5–17.0 cm in length, under surfaces densely whitehairy. Rare in mesic forest at an elevation of about 1,050 m on the Kōke‘e Plateau of Kaua‘i. When censused in 2003, there were 17 small populations with a total of not more than 114 individuals. Threats come from feral animals and alien plant species.

Remya mauiensis Hillebr. Pseudognaphalium sandwicensium var. molokaiense 4.94 

ENDEMIC | ENDANGERED

Erect to prostrate perennial herbs, up to 65 cm tall. Leaves moderately to very densely white-woolly pubescent, linear spatulate to spatulate, 1.0–6.5 cm in length. Flower heads in terminal leafless clusters. Occurs in dry sites from dunes near the coast to lava and cinder fields at elevations up to 3,000 m. The range of the species is from Kure and Midway atolls to all of the main islands except Kaho‘olawe. Four distinct varieties are recognized based on differences in leaf shape, pubescence, and the inflorescence, with the widespread var. sandwicensium pictured. Listed in the Manual as Gnaphalium sandwicensium var. molokaiense.

Remya—An endemic Hawaiian genus

of 3 shrub species, all of them rare and endangered. The name honors the French botanist Jules Remy, who collected in the Hawaiian Islands during the mid-nineteenth century. The affinities of this genus appear to be close to those of Olearia in the Southern Hemisphere. In addition to the 2 species pictured here, a third species (R. montgomeryi) is rare and endangered, with a highly restricted occurrence on Kaua‘i.

Leaves chartaceous, narrowly elliptic, 9–18 cm in length, densely grayish white pubescent on lower surface. Rare in mesic forest at elevations of 850–920 m on West Maui. There are 6 known populations with about 320 individuals. Threats come from alien plant species and fire.

Tetramolopium—A genus of about 38 species found in the Hawaiian Islands, the Cook Islands, Australia, and New Guinea. These range in growth form from prostrate herbs to sprawling or erect shrubs distributed in the Hawaiian Islands from sea level to above 3,000 m elevation. Eleven species, 10 endemic and 1 indigenous, are recognized from the Hawaiian Islands, with many of the taxa considered rare and endangered. Monecious or gynomonecious. The name is from the Greek, tetra, four, and molopium, stripe, because of the nerves on the achenes which vary from 1 to 4.

Tetramolopium arenarium (A. Gray) Hillebr. ENDEMIC | ENDANGERED/EXTINCT

4.96 

Remya mauiensis—habit

4.98 

Remya kauaiensis Hillebr.

Tetramolopium arenarium subsp. arenarium

ENDEMIC | ENDANGERED

4.97 

4.95 

Remya kauaiensis

Remya mauiensis

Sprawling, multibranched shrubs. 1–2 m tall, with ascending branches forming loosely tangled or sprawling clumps.

Erect shrubs to 3 m tall, stems pubescent or glandular pubescent when young. Leaves oblanceolate to oblong-lanceolate, 1.5–4.0 cm in length, glandular. Heads 5–11, corollas maroon. Formerly occurring in dry forest to wet forest at elevations of 800–1,500 on Maui and Hawai‘i. There are 2 subspecies. Pictured is subsp. arenarium native to East Maui (now extinct) and Hawai‘i (Hulalālai, Nohonaohae, Waimea). Once thought to

FLOWERING PLANTS

be extinct, the species was rediscovered in dry forest in the Pōhakuloa area on Hawai‘i in 1992. Today there are 5 known populations containing fewer than 500 individuals, with threats from alien plant species and fire. The subsp. laxum, with a more open inflorescence, once occurred on East Maui but is thought to be extinct.

Tetramolopium consanguineum (A. Gray) Hillebr. ENDEMIC | RARE/EXTINCT

Dwarf, caespitose shrubs to 20 cm tall with densely glandular stems. Leaves linear-oblanceolate to spatulate, 1.0–2.5 cm in length, densely glandular. Gynomonoecious, heads solitary or 5–7 per inflorescence, 20–30 flowers per head, rays white or tinged purple, corollas pink to purple. Occurs in subalpine woodland and alpine desert in crevices of lava flows or in cinder cones at elevations of 1,900– 3,300 m elevation on East Maui and Hawai‘i. Pictured is subsp. haleakalae from East Maui, while the subsp. humile occurs on Hawai‘i (Mauna Loa, Mauna Kea, Hualālai, Kīlauea).

Prostrate shrubs forming compact mats 5–10 cm tall and 8–40 cm in diameter. Leaves spathulate, 1.5–3.0 cm in length, densely pubescent or glandular pubescent. Monoecious, heads solitary, rays white, corollas yellow. Known only from lithified calcareous dunes in coastal shrubland at elevations of 10–200 m in the Mo‘omomi dune area of northwestern Moloka‘i. There are 2 named varieties, with var. rockii pictured. Threats today come from feral animals, alien plant species, and off-road vehicles.

Tetramolopium sylvae Lowrey INDIGENOUS | RARE

Tetramolopium lepidotum (Less.) Sherff ENDEMIC | ENDANGERED/EXTINCT

Tetramolopium consanguineum subsp. leptophyllum 4.99 

Erect shrubs 0.5–2.0 m tall, young stems with short glandular hairs. Leaves linear-oblanceolate, 1–6 cm in length, glandular. Gynomonoecious, heads 9–40, rays white or pink. Occurs in mesic forest at elevations of 200–1,600 m. Pictured is subsp. leptophyllum from the saddle area between Mauna Loa and Mauna Kea, Hawai‘i. The subsp. consanguineum from Kaua‘i, with shorter leaves, is presumed extinct.

Tetramolopium humile (A. Gray) Hillebr. ENDEMIC | SECURE

4.101 

Tetramolopium lepidotum subsp. lepidotum

Erect shrubs to 36 cm tall. Leaves filiform to linear or oblanceolate, 2.5–4.5 cm in length, glandular. Gynomonoecious, heads 6–12, flowers 20–40 per head, rays white to pinkish lavender, corollas maroon to pale salmon. Pictured is subsp. lepidotum from grassy ridgetops and west-facing cliffs at elevations of 600–900 m in the Wai‘anae Mountains, O‘ahu, and formerly Lāna‘i. Currently known from 3 populations and about 65 individuals. Threats come from feral animals and alien plant species. The subsp. arbusculum from Haleakalā, East Maui, has filiform leaves and is now considered to be extinct.

Tetramolopium rockii Sherff ENDEMIC | VULNERABLE

4.103 

Tetramolopium sylvae

Prostrate-rosette shrubs 4.5–6.0 cm tall, stems stout, branched at their base. Leaves succulent, spathulate, 1.5–3.0 cm in length, glabrous to slightly glandular. Monoecious, heads solitary, rays white, corollas yellow. Known from volcanic seacliffs along the windward coast of Moloka‘i and northwestern Maui. It has also been collected on calcareous beach deposits on Mitiaro in the Cook Islands.

Wilkesia—An endemic Hawaiian genus of

4.100 

Tetramolopium humile subsp. haleakalae

4.102 

Tetramolopium rockii var. rockii

2 species confined to the island of Kaua‘i. This is the third member of the famous silversword alliance that includes Argyroxiphium and Dubautia. Named after Charles Wilkes, who was the commander of the United States Exploring Expedition in the nineteenth century. The inflorescences of both species are highly glandular.

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FLOWERING PLANTS

Wilkesia gymnoxiphium A. Gray iliau

ENDEMIC | SECURE

Erect shrub to 70 cm tall, freely branched from near the base. Leaves grasslike, in whorls of 7–9, 10–20 cm in length. Occurs in a small population of a few hundred individuals on almost vertical dry ridges at 275–400 m elevation at Kā‘aweiki on the west end of Kaua‘i. Threats come from grazing by goats and fire.

Begoniaceae

(BEGONIA FAMILY)

4.104 

4.105 

Wilkesia gymnoxiphium—habit

Wilkesia gymnoxiphium

Erect, unbranched (rarely branched) rosette shrubs 1–5 m tall that usually die after flowering. Leaves in whorls of 9–15, 15–50 cm in length, joined to form a sticky-glandular sheathing cylinder. Inflorescence strongly glandular. Occurs on dry ridges or in open sites in dry to mesic forest at elevations of 425–1,100 m along Waimea Canyon and vicinity, Kaua‘i.

Mostly succulent herbs or shrubs, comprising about 1,400 species distributed in tropical lowlands and occasionally tropical uplands. Key features are the unisexual flowers, with the male flowers being zygomorphic and the ovaries usually angled or winged on the female flowers. Two genera are recognized, with Hillebrandia as a monotypic genus confined to the Hawaiian Islands. Many species of Begonia are cultivated as ornamentals. As many as 5–6 introduced species have become naturalized in the Islands.

Hillebrandia—A succulent perennial herb forming a monotypic genus endemic to the Hawaiian Islands. Named after William Hillebrand, who wrote the first flora of the Hawaiian Islands in 1888. Rather local in distribution and occasionally cultivated.

Hillebrandia sandwicensis Oliv. pua maka nui

ENDEMIC | VULNERABLE

Wilkesia hobdyi H. St. John dwarf iliau

ENDEMIC | ENDANGERED

4.106 

Wilkesia hobdyi

4.107 

Hillebrandia sandwicensis—habit

Erect perennial herbs 0.5–1.5 m tall. Leaves suborbicular, asymmetrical, 10–­ 25 cm in diameter, covered with short white papillate hairs. Flowers pink or white in a nodding inflorescence. Dies back during the winter. In open or shady wet ravines in wet forest at elevations of 600–1,850 m on Kaua‘i, Moloka‘i, Maui, and one old report from Mount Ka‘ala on O‘ahu. Hillebrandia is thought to represent the oldest plant lineage in the Hawaiian Islands, with molecular data suggesting that its ancestors evolved as much as 50 million years ago on the older and now submerged islands stretching past the existing Northwestern Hawaiian Islands.

Boraginaceae

(BORAGE FAMILY)

A large cosmopolitan family of trees, shrubs, herbs, and occasionally lianas. Key features are the usually coiled inflorescence and a dry fruit composed of 4 nutlets, although a drupe is characteristic of some genera. Foliage typically with one-celled bristly hairs. With the Hydrophyllaceae now included within the family, there are 142 genera and some 2,450 species. In the Hawaiian Islands represented by the indigenous genera Cordia, Nama, and Heliotropium. Six other genera have become naturalized, including the tree heliotrope (Tournefortia argentea). A family of little economic importance, except that a few genera are cultivated as ornamentals: forget-me-not (Myosotis), bluebells (Mertensia), heliotrope (Heliotropium), and Phacelia.

Cordia—A genus of about 250–300 species of woody shrubs and trees widely distributed in tropical and warm temperate regions around the world. Many species have fragrant, showy flowers and are popular in gardens, while other species are grown in plantations for their wood. A number of the species have edible fruit, with names that include clammy cherries, glue berries, sebesten, or snotty gobbles. The name honors father and son Euricius and Valerius Cordus, German botanists and pharmacists. In the Hawaiian Islands represented by 1 indigenous and 1 naturalized species and many cultivated species.

FLOWERING PLANTS

Cordia subcordata Lam. kou

INDIGENOUS | SECURE

Prostrate shrubs, sometimes forming mats. Stems usually decumbent to prostrate with silky leaves densely clustered toward the tips. Widely distributed from the Hawaiian Islands to Polynesia; occurs in sandy sites along the coast on Ni‘ihau, Kaua‘i, O‘ahu, Moloka‘i, rare on Maui and Hawai‘i, and possibly formerly on Lāna‘i and Kaho‘olawe. This is the endemic var. argenteum.

Nama sandwicensis A. Gray hinahina kahahai

ENDEMIC | VULNERABLE

Heliotropium curassavicum L.

kīpūkai, nena, seaside heliotrope 4.108 

Cordia subcordata

INDIGENOUS | SECURE

Brassicaceae

(MUSTARD FAMILY)

Heliotropium—A genus of about 350 spe-

Heliotropium anomalum Hook. & Arn. hinahina, pōhinahina

INDIGENOUS | SECURE

4.109 

Heliotropium anomalum

Nama sandwicensis

Annual or short-lived perennial herbs often forming mats up to 20 cm in diameter. The small thick leaves have inrolled margins. Flowers are blue or white. Scattered and uncommon in sandy soils or on raised limestone soil in strand and coastal vegetation at elevations from sea level up to 220 m on all of the main islands except Kaho‘olawe. Also on Lisianski and Laysan, where the flower color is white rather than purplish blue. Relationships are with species in the southwestern United States.

Small erect trees, readily producing aerial shoots from extensive shallow roots, sometimes forming thickets. Leaves thin, coriaceous, broadly ovate to elliptic, 10–120 cm in length. Flowers orange, fruit brown and buoyant when mature. Clearly brought to the Hawaiian Islands by the Polynesian settlers because of its high-quality wood used for bowls and utensils. Although a Polynesian introduction, it is now considered to be indigenous due to the discovery of fruit in soil layers that predates the Polynesian settlement. Widespread at low elevations in dry coastal areas of all of the main islands except Kaho‘olawe. cies of herbs and shrubs from warm temperate and tropical areas. The name is from the Greek helios, sun, and trope, turning, because it was believed that the flowers turned to face the sun, which in fact they do not. In the Hawaiian Islands there are 2 indigenous species and 2 that have become naturalized. Some species are grown for their showy flowers.

4.111 

4.110 

Heliotropium curassavicum

Glabrous perennial herbs with prostrate to decumbent, succulent stems to 45 cm in length. Leaves thick and succulent. Widely distributed in North America, South America, Australia, and beyond. In the Hawaiian Islands in wet or dry sites in coastal areas on Laysan, French Frigate Shoals, Nīhoa, and all of the main islands.

Nama—A genus of 45 species of annual or perennial pubescent herbs of the southwestern United States and Mexico, South America, and the Hawaiian Islands. The name is from the Greek word for spring. In the Hawaiian Islands there is 1 endemic species found in coastal habitats.

A large cosmopolitan family of mostly herbs but occasionally shrubs. Key features are the presence of 4 sepals and 4 petals diagonally disposed, 6 stamens, of which 2 are short and 4 long arranged in 2 whorls, and the unique fruit called a silicle or silique. There are 321 genera and about 3,400 species. Economically the family is important for food crops such as cabbage, cauliflower, broccoli, turnips (all Brassica), radishes (Raphanus), and watercress (Rorippa). It is also the source of condiments such as mustard (Brassica) and horseradish (Armoracia). Cultivated plants in the family include alyssum (Lobularia) and candytuft (Iberia). In the Hawaiian Islands represented by the indigenous genus Lepidium. Weedy genera such as Brassica, Sisymbrium, Cardamine, Capsella, and Descurainia are naturalized.

Lepidium—A genus of about 220 species

of cosmopolitan herbs, occasionally subshrubs or shrubs that are known as pepperworts. The name is from the Greek lepis, scale, which refers to the scalelike form of the small flat fruit. Some species are cultivated and eaten in salads. In the

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FLOWERING PLANTS

Hawaiian Islands there are 4 endemic and 1 indigenous species. These form a clade that is sister to a group of herbaceous species from the southwestern United States and Mexico.

Lepidium bidentatum Montin

kūnānā, naunau, ‘ānaunau, ‘ānounou INDIGENOUS | VULNERABLE

4.114 

Lepidium serra

Small, many-branched shrubs to 1.2 m tall with elliptic leaves crowded at the ends of the branches. Rare on rocky ledges and cliff faces in mesic forest at elevations of 610–1,160 m on Kaua‘i.

4.112 

Lepidium bidentatum var. o-waihiense

Subshrubs up to 60 cm tall with ascending to weakly erect stems. The leaves are thick and somewhat fleshy. Occurs in coastal sites and low elevation from sea level to 240 m on dry and rocky slopes on Kure, Midway, and Pearl and Hermes atolls, Laysan (extinct), and on all of the main islands except Ni‘ihau and Kaho‘olawe. Hawaiian plants represent the endemic var. o-waihiense, representing a species widespread beyond the Hawaiian Islands.

Lepidium serra H. Mann ‘ānaunau, peppergrass ENDEMIC | SECURE

4.113 

Lepidium serra—habit

Rorippa—A widespread genus of annual to biennial aquatic herbs with about 85 species. The genus has a north temperate center of distribution, ranging from Europe and North Africa across central Asia to North America. The name comes from an Old Saxon word for these plants and perhaps other aquatic species. In the Hawaiian Islands represented by Rorippa sarmentosa, which occurs in wet, sometimes disturbed areas on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i. Perhaps indigenous but often considered to be a deliberate or unintentional Polynesian introduction. Listed in the Manual as Nasturtium sarmentosum. Not pictured.

separated on their architecture, growth habit, and structure of fruit. Clermontia species are typically branched shrubs or small trees up to 7 m tall, with typically 1–3 flowers per inflorescence and fleshy fruit. Cyanea and Delissea species are typically unbranched or branching only at the base, with a cluster of relatively broad leaves at the apex, 5 or more flowers in the inflorescence, and fleshy fruits. Delissea differs from Cyanea in the presence of a small dorsal knob on the corolla. Lobelia and Trematolobelia have long, thin leaves growing from a single semiwoody stem and capsular fruit with wind-dispersed seeds. The inflorescence is unbranched in Lobelia and branched in Trematolobelia. Brighamia species are distinctive plants with a short, thick stem topped by a dense cluster of broad leaves, elongate white or yellow flowers, and capsular fruit. Economically the family is important for many ornamentals, such as harebells (Campanula), Lobelia, and Platycodon. Hippobroma and Wahlenbergia are introduced genera.

Brighamia—An endemic Hawaiian genus of 2 species, both of them quite rare. The name honors W. T. Brigham, who was the first director of the Bishop Museum and who studied the native flora. The succulent nature of the stem and the showy white or yellow flowers make this a desirable ornamental.

Brighamia insignis A. Gray

‘ōlulu, pū aupaka

ENDEMIC | ENDANGERED

Campanulaceae

(BELLFLOWER FAMILY)

A family of herbs, shrubs, or trees, with some members possessing a network of latex-bearing glands. Key features are an inferior ovary and the stamens frequently united by coherence of the anthers. The subfamily Lobelioideae, to which the Hawaiian genera belong, is characterized by irregular corollas and joined anthers. There are 79 genera and about 1,900 species worldwide. In the Hawaiian Islands, besides the indigenous genus Lobelia, there are also 5 endemic genera: Brighamia, Clermontia, Cyanea, Delissea, and Trematolobelia. These 5 genera and their 141 species represent an evolutionary lineage from a single colonizing taxon. The 5 genera can be broadly

4.115 

Brighamia insignis

Unbranched stem succulents with tapering stem 1–5 m tall. Leaves fleshy, 12–15 cm in length, arranged in a terminal rosette. Corolla yellow, calyx lobes 0.1– 1.0 mm in length, pedicels 10–30 mm in length. Occurs on sea cliffs at elevations from sea level to 470 m on Kaua‘i (Nāpali Coast, Hā‘upu Ridge, Nāwiliwili), and

FLOWERING PLANTS

previously known from Ni‘ihau (Ka‘ali Cliff) (see figure 2.1). There are 5 extant populations totaling about 60–70 plants, but they are widely cultivated. Threats come from goats and alien plant species.

Clermontia arborescens (H. Mann) Hillebr.

Clermontia clermontioides (Gaudich.) A. Heller

ENDEMIC | SECURE/RARE

ENDEMIC | SECURE

‘ōhā wai, hāhā, ‘ōhā wai nui

‘ōhā wai, hāhā, ‘ōhā

Brighamia rockii H. St. John pua ‘ala

ENDEMIC | ENDANGERED

4.117 

4.116 

Brighamia rockii

Unbranched stem succulents with tapering stem 1–5 m tall. Leaves fleshy, 6–22 cm in length arranged in a terminal rosette. Corolla white, calyx lobes 2.5–8.0 mm in length, pedicels 6–12 mm. Occurs on sea cliffs at elevation up to 470 m on windward Moloka‘i (Kalaupapa to Hālawa), Lāna‘i (Maunalei Valley, extinct), and Maui (extinct). The 5 known populations total fewer than 200 individuals, with threats from feral animals.

Clermontia—An endemic genus of 22 species of branched shrubs or small trees (occasionally epiphytic) with milky sap and fleshy fruit. These are never palmlike in form. The name honors M. le Marquis de Clermont-Tonnerre, who was minister of the French Navy during the Freycinet expedition in the early nineteenth century. These plants are usually found in wet forest or along the margins of bogs and tend to occupy more open places or clearings in the forest. The curved flowers are (or were) pollinated by native birds such as the honeycreepers. Flower color ranges from black-purple to violet, magenta, green, and white.

Clermontia arborescens var. waihiae

Terrestrial or epiphytic trees or shrubs to 8 m tall. Leaves glossy dark green above, greenish white below; elliptic to oblong with blades 10–23 cm in length. Inflorescence 1–2 flowered, corolla fleshy greenish white and among the largest in the genus with a length of 5.5–8.5 cm. Occurs in mesic to wet forest at elevations of 520– 1,825 m. on Moloka‘i, Lāna‘i, and Maui. Pictured is subsp. waihiae from Maui (see figure 2.1), which has shorter calyx lobes than the rare subsp. arborescens on West Maui (‘Īao Valley and Hana‘ula). The subsp. waikoluensis occurs on Moloka‘i and Lāna‘i.

4.119 

Clermontia clermontioides subsp. rockiana

Clermontia calophylla E. Wimm.

Terrestrial or epiphytic shrubs or trees to 6.5 m tall. The flowers in this species may be either green, white, or purplish. Leaves green above, pale green below, oblong, elliptic or oblanceolate, blades 7–25 cm in length. Inflorescence 1–2(–4) flowered, corolla greenish, white, or purplish. Occurs in mesic to wet forest at elevations of 670–1,825 m on Hawai‘i. Two subspecies are recognized based on the length of the calyx lobes. Pictured is subsp. rockiana, which occurs in the Kona District (Mauna Loa and Hualālai, Kealia to Pu‘u Wa‘awa‘a). The subsp. clermontioides occurs in the same area from Pāhala to Pu‘u Lehua.

ENDEMIC | RARE

Clermontia drepanomorpha Rock

‘ōhā wai, hāhā

‘ōhā wai, hāhā

ENDEMIC | ENDANGERED

4.118 

Clermontia calophylla

Terrestrial shrubs 1.5–5.0 m tall. Leaves green above, pale below, broadly elliptic to elliptic, blades 12–19 cm in length. Inflorescence 2(–4) flowered; similar in appearance to C. parviflora but with a larger dark purple flower and larger leaves and berries. Occurs in wet forest at 885– 1,460 m elevation in the Kohala Mountains, Hawai‘i. The species is considered to be critically endangered and known from only 3 small populations with an unknown number of individuals. Threats come from feral pigs and alien plant species.

4.120 

Clermontia drepanomorpha

Terrestrial or epiphytic trees, 2–7 m tall. Leaves green above, pale green or purplish below, oblanceolate to elliptic, 8–22 cm in length. Inflorescence 2(–4) flowered; flowers blackish purple. Occurs in wet boggy forest at elevations of 915– 1,460 m on the summit of the Kohala Mountains, Hawai‘i. Existing populations number less than 250 individuals, with threats from feral animals and invasive plant species.

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FLOWERING PLANTS

Clermontia fauriei H. Lév.

hāhā‘aiakamanu, ‘ōhā wai, hāhā ENDEMIC | SECURE

in wet forest and along the margins of bogs at 530–1,975 m elevation on Moloka‘i, Lāna‘i, and Maui. There are 3 named subspecies, with the subsp. grandiflora from West Maui. The subsp. maxima is restricted to East Maui and endangered, while the subsp. munroi occurs on Moloka‘i, Lāna‘i, and Maui.

Clermontia hawaiiensis (Hillebr.) Rock

Terrestrial or epiphytic shrubs or trees, 1–6 m tall. Leaves green above, pale green below, elliptic, oblanceolate or narrowly obovate, blades 12–26 cm in length. Inflorescence 2(–7) flowered, perianth pale green externally, greenish white inside. Occurs in mesic forest or on the margins of wet forest at elevations of 245–1,100 m on O‘ahu, Moloka‘i, and Maui. A common species at lower elevations.

ENDEMIC | SECURE

Clermontia kohalae Rock

‘ōhā wai, hāhā

‘ōhā wai, hāhā 4.121 

ENDEMIC | SECURE

Clermontia fauriei

Terrestrial or epiphytic trees or shrubs, 2–7 m tall. Leaves dark green and glossy above, pale green below, oblong to elliptic, blades 5–17 cm in length. Inflorescence 1–2 flowered, corolla strongly curved, 6–7 cm in length, greenish or purplish externally and cream-colored internally. Occurs in diverse mesic to wet forest at elevations of 365–1,400 m on Kaua‘i and O‘ahu and is the only member of the genus present on the former island.

4.125 

Clermontia grandiflora Gaudich. ‘ōhā wai, hāhā

ENDEMIC | SECURE/ENDANGERED

4.123 

Clermontia hawaiiensis

Terrestrial or epiphytic trees or shrubs, 1–9 m tall. Leaves dark green above, whitish green below, oblong to oblanceolate, blades 9–24 cm in length. Inflorescence 2(–4) flowered, perianth greenish white sometimes tinged at the apex with purple, curved, 5.0–6.5 cm in length. Occurs in mesic to wet forest at 550–1,760 m elevation in the Puna and Ka‘ū districts, Hawai‘i.

Clermontia kohalae

Terrrestial or ephiphytic trees, 2–6 m tall. Leaves narrowly elliptic to oblanceolate, blades 10–21 cm in length. Inflorescence 2(–4) flowered, perianth blackish purple. Occurs in wet forest at elevations of 370– 1,370 m in the Kohala, Hāmākua, and North Kona districts of Hawai‘i.

Clermontia lindseyana Rock ‘ōhā wai, hāhā

ENDEMIC | ENDANGERED

Clermontia kakeana Meyen ‘ōhā wai, hāhā

ENDEMIC | SECURE

4.122 

4.126 

Clermontia grandiflora subsp. grandiflora

Terrestrial trees, 1–6 m tall, rarely epiphytic. Leaves green above, pale green below, elliptic to oblanceolate, blades 4–15 cm in length. Inflorescence 2(–5) flowered, perianth curved 5.5–8.0 cm in length, pendulous, corolla greenish to purplish or rose colored. Occurs

4.124 

Clermontia kakeana

Clermontia lindseyana

Terrestrial shrubs or trees, 2.5–6.0 m tall. Leaves dark green above, pale green to purplish below, oblanceolate to elliptic, blades 13–24 cm in length. Inflorescence 2 flowered, perianth green, sometimes tinged with purple on the outside, white or cream inside. Rare in mesic

FLOWERING PLANTS

forest at 1,220–1,825 m elevation on East Maui (leeward slopes of Haleakalā) and Hawai‘i (South Hilo, North Hilo, Ka‘ū, and South Kona districts). The 10 known populations total fewer than 1,000 individuals, with threats from feral animals, deforestation, and alien plant species.

Clermontia montis-loa Rock ‘ōhā wai, hāhā

ENDEMIC | SECURE

Occurs in mesic valleys to wet forest at 400–1,200 m elevation on O‘ahu, Moloka‘i, Lāna‘i, and Maui. There are 3 named subspecies, with the subsp. oblongifolia from O‘ahu pictured. Both the subsp. brevipes restricted to Moloka‘i and the subsp. mauiensis from Lāna‘i (extinct) and Maui are endangered.

Clermontia parviflora Gaudich. ex A. Gray ‘ōhā wai, hāhā

ENDEMIC | SECURE

Clermontia pallida Hillebr. ‘ōhā wai, hāhā

ENDEMIC | SECURE

4.131 

4.127 

Terrestrial or epiphytic shrubs 1.0–3.5 m tall. Leaves green above, pale green below, elliptic to oblanceolate, blades 6–18 cm in length. Inflorescence 3–7 flowered, perianth green, white or purple externally, white or pale purple inside. Occurs in wet forest at elevations of 120–1,460 m in the Kohala Mountains and on the windward slopes of Mauna Kea and Mauna Loa, Hawai‘i. Hybridizes with other species of Clermontia where they grow together.

Clermontia montis-loa

Terrestrial or epiphytic trees or shrubs, 2–5 m tall. Leaves dark green above, pale green to purplish below, pubescent, oblanceolate to oblong, blades 8–19 cm in length. Inflorescence 2–3 flowered, perianth greenish to purple externally, purple to dark purple inside. Occurs in wet forest at elevations of 1,070–1,700 m on the eastern side of Hawai‘i (North Hilo, South Hilo, Puna, and Ka‘ū districts). This species hybridizes with C. parviflora, the intermediates being variable in color from magenta to white.

Clermontia parviflora

Clermontia peleana Rock ‘ōhā wai, hāhā

ENDEMIC | ENDANGERED

4.129 

Clermontia pallida—habit

4.130 

Clermontia pallida

Clermontia oblongifolia Gaudich. ‘ōhā wai, hāhā

ENDEMIC | SECURE

4.128 

Clermontia oblongifolia subsp. oblongifolia

Terrestrial shrubs or trees, 2–7 m tall. Leaves dark green and glossy above, whitish green below, oblong to lanceolate, blades 7–19 cm in length. Inflorescence 2(–3) flowered, perianth greenish white to purple externally, white or cream inside.

Terrestrial or rarely epiphytic shrubs or trees 2–5 m tall. Leaves green above, whitish green below, oblong to elliptic or oblanceolate, blades 7–15 cm in length, petioles yellowish white and translucent. Inflorescences 2–3(–7) flowered, perianth blackish purple to dark maroon externally, pale greenish inside. Occurs in wet forest at elevations of 915–1,390 m on Moloka‘i.

4.132 

Clermontia peleana subsp. peleana

Epiphytic shrubs or trees 1.5–6.0 m tall. Leaves oblong to elliptic, blades 8–20 cm in length. Inflorescence 1–2 flowered, corolla curved, white (on East Maui) or dark purple (on Hawai‘i). Very rare and thought to be extinct until its recent rediscovery. Once present in wet forest at elevations of 530–1,150 m on windward slopes of East Maui and Hawai‘i (Kohala Mountains, Mauna Kea, and

75

76

FLOWERING PLANTS

Mauna Loa). There are 2 named subspecies, both endangered. The subsp. peleana once occurred on the windward slopes of Mauna Kea and Mauna Loa but is now known only from cultivation. The subsp. singuliflora once occurred on the windward slopes of East Maui and Mauna Kea and was considered to be extinct until a few plants were discovered in 2010 in the Kohala Mountains.

Clermontia persicifolia Gaudich. ‘ōhā wai, hāhā

ENDEMIC | SECURE

Terrestrial shrubs or trees, 4–5 m tall. Leaves oblanceolate to obovate, blades 9–20 cm in length. Inflorescence 1–2 flowered, corolla rose colored externally. Restricted in distribution to wet forests, typically along streambeds, at elevations of 1,650–1,825 m on East Maui. This species is closely related to C. arborescens but differs in the rose color of the perianth and tuberculate surface of the base of the floral tube. Population size is uncertain, but there are no more than a few thousand plants. Threats come from feral pigs and alien plant species.

Clermontia waimeae Rock ‘ōhā wai, hāhā

ENDEMIC | SECURE

Clermontia with usually few flowered inflorescences, those of Cyanaea are many flowered. Additionally, the plants are often unbranched and palmlike in growth form. Some species are armed with prickles. Many have juvenile leaves that are quite different from the adults. The smallest species is less than 1 m tall, while the tallest may be up to 14 m tall. The name is from the Greek cyaneos, blue, supposedly because the type species had blue flowers. Eight of these species, restricted to O‘ahu with one exception, were formerly segregated as the genus Rollandia. Many new species have been described in recent years. From 60 described species of Cyanea in the Manual, including Rollandia, the genus now has an added 19 species.

Cyanea aculeatiflora Rock hāhā

ENDEMIC | SECURE

4.133 

Clermontia persicifolia

Terrestrial or epiphytic shrubs or trees, 2–6 m tall. Leaves dark glossy green above, pale green below, elliptic to oblanceolate, blades 7–16 cm in length. Inflorescence 2–3(–6) flowered, perianth gently curved, white to greenish white externally. Occurs in wet forests at elevations of 305–850 m in the Wai‘anae and Ko‘olau Mountians, O‘ahu, although less commonly in the Wai‘anae Mountains.

Clermontia tuberculata C. N. Forbes ‘ōhā wai, hāhā

ENDEMIC | RARE

4.134 

Clermontia tuberculata

4.135 

4.136 

Cyanea aculeatiflora—habit

4.137 

Cyanea aculeatiflora

Clermontia waimeae

Terrestrial or epiphytic shrubs or trees, 1–7 m tall. Leaves oblong to elliptic or oblanceolate, blades 5–15 cm. Inflorescence 2–3(–5) flowered, corolla blackish purple. Occurs in bogs and wet forest at 1,070–1,520 m elevation in the Kohala Mountains, Hawai‘i. Known from only a few localities with no more than a few thousand plants. Threats come from alien plant species and feral pigs.

Cyanea—An endemic genus now including 79 endemic species of branched or unbranched shrubs or trees with white, tan, or yellow latex. These are plants of wet forest or occasionally mesic forest habitats, with several species becoming palmlike in growth form. Unlike

Palmlike trees to 7 m tall. Leaves in an apical rosette, prickly on the veins and petioles; oblanceolate, oblong or elliptic, blades 40–100 cm in length. Inflorescence 5–10 flowered; corolla blackish purple, prickly externally. Uncommon in wet forest at elevations of 945–1,450 m on the windward slopes of Haleakalā, East Maui.

FLOWERING PLANTS

Cyanea acuminata (Gaudich.) Hillebr. hāhā

ENDEMIC | RARE

Ko‘olau Mountains and less frequently in the Wai‘anae Mountains, O‘ahu. Also occurs in mesic forest on Moloka‘i, Lāna‘i, and West Maui. Estimates suggest a population of up to 10,000 plants.

Cyanea arborea Hillebr. hāhā

ENDEMIC | EXTINCT

Palmlike trees 4–8 m tall. Leaves oblanceolate, blades 65–90 cm in length. Inflorescence 15–25 flowered, corolla cream tinged with pale lilac. Formerly occurring in mesic forest at 1,520–1,650 m elevation on the leeward slopes of Haleakalā, Maui. Last collected in 1928. 4.138 

Cyanea acuminata

Shrubs 0.3–2.0 m tall, stems woody only at their base, typically unbranched. Leaves lanceolate to narrowly oblong, blades 9–32 cm in length. Inflorescence 6–25 flowered, corolla white to greenish white, sometimes tinged with purple. Occurs in mesic valleys at elevations of 300–730 m in the Ko‘olau Mountains, O‘ahu. Currently there are 12 known populations with a toal of 149–175 individuals. Threats come from feral animals, invasive plant species, and fire.

Cyanea angustifolia (Cham.) Hillebr. hāhā

ENDEMIC | SECURE

Cyanea asarifolia H. St. John hāhā

4.140 

hāhā

ENDEMIC | VULNERABLE/EXTINCT

Cyanea asarifolia

Low shrub 0.3–1.0 m tall, sparingly branched. Leaves heart shaped and cordate at their base, blades 8.5–10.5 cm in length, 7–8 cm in width. Inflorescence 30–40 flowered, corolla white with longitudinal purple stripes. Rare on sheer rocky cliffs at an elevation of about 330 m above the bed of Anahola Stream, northeast Kaua‘i. Known from a single population of 20–30 individuals, with threats from landslides, feral pigs, alien plant species, and slugs.

hāhā

ENDEMIC | RARE

Cyanea angustifolia

Branched shrubs, 1.5–5.0 m tall. Leaves lanceolate to narrowly elliptic or oblong, blades 9–32 cm in length. Inflorescences 6–25 flowered; corolla white to greenish white. A common species in mesic to wet forests at elevations of 150–750 m in the

Cyanea copelandii Rock

ENDEMIC | ENDANGERED

Cyanea calycina (Cham.) Lammers

4.139 

Trailing shrubs, stems 1–3 m in length, stems rough with sharp points in juveniles. Leaves elliptic to oblanceolate, blades 15–60 cm in length, lower leaf surface densely pubescent with branched and clustered hairs. Inflorescence 4–16 flowered, corolla pale to dark magenta. Occurs in mesic valleys and wet forest at elevations of 300–915 m in the Wai‘anae and Ko‘olau Mountains, O‘ahu. Closely resembles Cyanea lanceolata. Known from 26 populations with less than 500 plants. Threats come from feral pigs and goats. Listed in the Manual as Rollandia lanceolata subsp. calycina.

4.141 

Cyanea calycina

4.142 

Cyanea copelandii subsp. haleakalaensis

Trailing liana-like shrubs 0.3–2.0 m tall with tan latex and semiherbaceous stems, often reclining on moss-covered tree trunks. Leaves oblanceolate to elliptic, blades 10–27 cm in length. Inflorescence 5–12 flowered, corolla yellowish but appears pale rose due to the dark red pubescence. The subsp. copelandii, with narrower leaves, once occurred on the windward slopes of Mauna Loa, Hawai‘i, and is now considered to be extinct. The subsp. haleakalensis pictured occurs in wet forest at elevations of 660–1,600 m on East Maui. Three small populations are known numbering less than 250 individuals. Threats come from feral grazing animals and alien plant species.

77

78

FLOWERING PLANTS

Cyanea coriacea (A. Gray) Hillebr.

Cyanea dolichopoda Lammers & Lorence

Cyanea elliptica (Rock) Lammers

ENDEMIC | SECURE

ENDEMIC | EXTINCT?

ENDEMIC | SECURE

hāhā

hāhā

4.145 

4.143 

Cyanea coriacea

Unbranched or lightly branched shrubs 2.5–3.0 m tall. Leaves oblong to oblanceolate, blades 20–38 cm in length. Inflorescence 12–20 flowered, white tinged with purple. Occurs in mesic valleys at 150–220 m elevation on northern Kaua‘i.

hāhā

Cyanea dolichopoda

Branched shrub up to 2 m tall. Leaves ovate to oblong, 4–6 cm in length and 2–3 cm in width. First discovered in 1990, with a few plants growing on wet cliffs at an elevation of about 700 m in the Blue Hole on the north fork of the Wailua River, Kaua‘i. Not seen since 1992 and possibly now extinct. Not included in the Manual.

Cyanea dunbariae Rock hāhā

ENDEMIC | ENDANGERED

Cyanea crispa (Gaudich.) Lammers, Givnish & Sytsma hāhā ENDEMIC | ENDANGERED

4.147 

Cyanea elliptica

Branched shrubs 1–4 m tall. Leaves elliptic to oblong, 8–18 cm in length, glabrous or pubescent along the midrib. Inflorescence 6–14 flowered, corolla greenish white or white tinged with purple. Occurs in mesic valleys and forests at elevations of 475–915 m on Lāna‘i and Maui. Estimates suggest a population of up to 10,000 plants.

Cyanea fissa (H. Mann.) Hillebr. hāhā

ENDEMIC | SECURE

4.144 

Cyanea crispa

Small shrub, stems fleshy, 0.3–1.3 m in length. Leaves broadly obovate, blades 30–75 cm in length. Inflorescence 3–8 flowered, corolla pale magenta with darker purple stripes. Occurs in mesic to wet forest at elevations of 185–730 m in the Ko‘olau Mountains, O‘ahu. Currently known from 7 populations with about 56 individuals. Threats come from feral animals, slugs, and alien plant species. Listed in the Manual as Rollandia crispa.

4.146 

Cyanea dunbariae

Branched shrub 1.5–2.0 m tall, stems smooth. Leaves ovate to broadly elliptic, blades 10–22 cm in length. Inflorescence 6–8 flowered, corolla white tinged with pale lilac. Rare in wet forest in Waihānau and Mokomoko gulches, Moloka‘i. In 2002 there were only 16 plants remaining in the wild. Threats come from feral animals, invasive alien plants, and slugs.

4.148 

Cyanea fissa

Unbranched or sparingly branched shrubs 1.5–5.0 m tall, with yellow latex. Leaves oblanceolate to obovate, blades 32–80 cm in length. Inflorescence 8–20 flowered, corolla white to pale lilac, striped with purple. Occurs in mesic and wet forests and bogs at elevations of 200– 915 m on Kaua‘i.

FLOWERING PLANTS

Low shrubs to 1–6 m tall, usually unbranched and woody only at the base. Leaves oblanceolate to oblong, 10–27 cm in length. Inflorescence 5–12 flowered, corolla sometimes tinged with purple. Occurs in wet forests at elevations of 1,030–1,400 m on Kaua‘i.

Cyanea floribunda E. Wimm. hāhā

ENDEMIC | SECURE

Cyanea horrida (Rock) O. Deg. & Hosaka 4.151 

4.149 

Cyanea floribunda

Low shrub 0.3–1.2 m tall, usually unbranched and woody only at the base, tan latex. Leaves elliptic, blades 15–27 cm long. Inflorescence 6–16 flowered, corolla white, sometimes tinged with purple. Occurs in wet forest at 460–1,520 m elevation in the Kohala Mountains and windward Mauna Kea and Mauna Loa, Hawai‘i. Listed in the Manual as Cyanea degeneriana.

Cyanea grimesiana Gaudich. hāhā

ENDEMIC | ENDANGERED

Cyanea grimesiana subsp. obatae

Shrubs from 1–3.2 m tall with branched or sparingly branched stems. A distinctive species because of its pinnatified leaves, blades 27–58 cm in length, lower surface with small spines along the midrib. Inflorescence 6–12 flowered, corolla purplish to greenish or yellowish white. Occurs in mesic to wet forest at 200–730 m on O‘ahu and Moloka‘i. Two subspecies are known, with both extremely endangered. Pictured is subsp. obatae from the southern Wai‘anae Mountains, O‘ahu. This taxon is known from 8 populations totaling 41 individuals. The subsp. grimesiana occurs in the Ko‘olau Mountains and on Mount Ka‘ala, O‘ahu, and on Moloka‘i. The Manual recognized an extinct subsp. cylindrocalyx from Waipi‘o Valley, Hawai‘i, but this taxon is now raised to species rank as Cyanea cylindrocalyx.

Palmlike trees 1–4 m tall. Leaves pinnately lobed or divided, especially in juveniles, blades 20–37 cm in length, leaf surface with sharp points. Inflorescence 5–8 flowered, clustered among the leaves, corolla blackish purple to greenish white tinged with purple. Occurs in wet forest and the margins of subalpine forest on the windward slopes of Haleakalā, East Maui (see figure 2.11). Known from only 12 populations and 44 individuals.

Cyanea humboldtiana (Gaudich.) Lammers, Givnish & Sytsma hāhā

ENDEMIC | ENDANGERED

hāhā

ENDEMIC | ENDANGERED

Palmlike trees, 3–8 m tall. Leaves oblanceolate to oblong, 50–80 cm in length. Inflorescence 5–10 flowered, corolla magenta. Rare in wet ‘ōhi‘a forest at elevations of 1,200–1,400 m on the windward slopes of Haleakalā (subsp. hamatiflora) (see figure 2.1) and 1,300–1,740 m in the Kona District, Hawai‘i (subsp. carlsonii).

hāhā

ENDEMIC | SECURE

Cyanea grimesiana subsp. obatae—habit

ENDEMIC | ENDANGERED

Cyanea hamatiflora Rock

Cyanea hirtella (H. Mann) Hillebr.

4.150 

hāhā

4.152 

Cyanea hirtella

4.153 

Cyanea humboldtiana

Unbranched shrubs, 1–2 m tall. Leaves obovate to broadly elliptic, blades 18–45 cm. Inflorescence 5–12 flowered, pendent, corolla dark magenta or rarely white, peduncles of the inflorescence may be up to 25 cm long. Occurs in wet forest at elevations of 300–915 m in the Ko‘olau Mountains, O‘ahu. Threats to remaining plants come from feral pigs and alien plant species. Listed in the Manual as Rollandia humboldtiana.

79

80

FLOWERING PLANTS

Cyanea koolauensis Lammers, Givnish & Sytsma hāhā

ENDEMIC | ENDANGERED

totaling a few hundred individuals and is declining in numbers. Threats come from feral pigs and goats, rats, and alien plant species.

Cyanea lanceolata (Gaudich.) Lammers, Givnish & Sytsma hāhā ENDEMIC | RARE

4.154 

Cyanea koolauensis

Shrubs with stems 1.0–1.5 m in length. Leaves linear to narrowly elliptic, blades 16–36 cm in length, lower leaf surface whitish. Inflorescence 3–6 flowered, corolla dark magenta. Occurs in cloudswept wet forest on ridgelines, less often in mesic valleys, in the Ko‘olau Mountains, O‘ahu. As of 2009 there were 10 scattered populations totaling approximately 160 individuals. Threats come from military training, feral animals, and alien plant species. Listed in the Manual as Rollandia angustifolia.

Cyanea kunthiana Hillebr. hāhā

ENDEMIC | RARE

4.158 

4.156 

Cyanea lanceolata

Unbranched shrubs, 1–3 m tall, juvenile stems prickly with sharp points. Leaves elliptic or oblanceolate, blades 15–60 cm in length, lower leaf surface pubescent with simple hairs, juvenile leaves prickly. Inflorescence 4–16 flowered, corolla pale to dark magenta, rarely pale greenish. Occurs in mesic valleys and wet forests 300–915 m in both the Wai‘anae and Ko‘olau Mountains, O‘ahu. Population estimates are fewer than 300 individuals. Threats come from feral pigs and alien plant species. Listed in the Manual as Rollandia lanceolata.

Cyanea leptostegia

Palmlike trees 3–14 m tall, with bright yellow latex. The leaves are dimorphic; juveniles are pinnately lobed with blades 50–70 cm in length, adult leaves linear to narrowly oblanceolate, blades 50–100 cm in length, margins undulate. Inflorescences 10–25 flowered, densely clustered among the leaves, corolla rose to purple. Occurs in diverse mesic forest at elevations of 1,000–1,300 m on western Kaua‘i. This is the tallest member of the genus. Still fairly common, with an estimated 7,000–10,000 mature individuals.

Cyanea lobata H. Mann hāhā

ENDEMIC | ENDANGERED

Cyanea leptostegia A. Gray hāhā lua, hāhā

ENDEMIC | VULNERABLE

4.159  4.155 

Cyanea kunthiana

Low shrubs to 0.5–1.5 m tall, usually unbranched and woody only at the base. Leaves elliptic to narrowly obovate, blades 12–30 cm long. Inflorescence 8–24 flowered, corolla whitish with lilac longitudinal stripes or dark purple. Occurs in wet forest at elevations of 1,000–1,600 m on Maui. Known from 15 populations

4.157 

Cyanea leptostegia—habit

Cyanea lobata subsp. lobata

Sparingly branched shrubs 1.3–2.3 m tall. Leaves oblong to ovate, blades 30–50 cm in length, upper surface glabrous, lower surface sparsely pubescent, leaf margins irregularly lobed. Inflorescence 5–12 flowered, greenish white to purplish. Rare in mesic valleys at elevations of 550– 900 m with subsp. baldwinii on Lana‘i (now possibly extinct) and subsp. lobata on northern West Maui.

FLOWERING PLANTS

Cyanea longiflora (Wawra) Lammers, Givnish & Sytsma hāhā

Cyanea mceldowneyi Rock

Cyanea pilosa A. Gray

ENDEMIC | ENDANGERED

ENDEMIC | SECURE

hāhā

hāhā

ENDEMIC | ENDANGERED

4.160 

Cyanea longiflora

4.162 

Cyanea mceldowneyi

Unbranched shrubs, 1–3 m tall, stems with small spines when juvenile. Leaves elliptic to oblanceolate, blades 30–55 cm in length. Inflorescence 5–10 flowered, corolla dark magenta. An uncommon species in mesic to wet forest at elevations of 275–760 m in the Wai‘anae and central Ko‘olau Mountains, O‘ahu. There are 3 populations totaling less than 300 individuals, with threats coming from feral animals, alien plant species, feral animals, military training, and vandalism. Listed in the Manual as Rollandia longiflora.

Unbranched shrubs 2–3 m tall, with prickly stems. Leaves are dimorphic: juveniles are prickly on both surfaces, while those of the adults are smooth or slightly prickly, adult leaves oblanceolate to narrowly oblong, blades 20–35 cm long. Inflorescence 5–7 flowered, corolla white with purple stripes. A rare plant in wet forest at elevations of 925–1,280 m on the windward slopes of Haleakalā, East Maui. In 2007 there were an estimated 60 plants in 2 populations. Threats come from feral animals and alien plant species.

Cyanea macrostegia Hillebr.

Cyanea membranacea Rock

ENDEMIC | SECURE

ENDEMIC | RARE

hāhā

hāhā

4.164 

Cyanea pilosa subsp. longipedunculata

Shrubs 0.8–2.0 m tall, woody only at the base, unbranched or sparingly branched at the base. Leaves broadly elliptic or obovate, pubescent on both surfaces but denser on the lower surface, blades 15–42 cm long. Inflorescence 6–28 flowered, pubescent, corolla white streaked with purple. One of the smallest members of the genus. Two subspecies are recognized based on the number of flowers in the inflorescence. Pictured is the subsp. longipedunculata from wet forest at elevations of 610–1,520 m on windward Mauna Kea and Mauna Loa, Hawai‘i. The subsp. pilosa occurs in similar habitats in the Kohala Mountains and windward Mauna Kea.

Cyanea pinnatifida (Cham.) E. Wimm. hāhā

ENDEMIC | ENDANGERED

4.163 

4.161 

Cyanea macrostegia

Palmlike trees 1–7 m tall. Leaves oblanceolate to elliptic, blades 20–80 cm in length. Inflorescence 5–15 flowered, clustered among the leaves, corolla blackish purple externally, white or pale lilac inside. Occurs in wet forests at 760–1,970 m elevation on Maui.

Cyanea membranacea

Sparingly branched shrubs 2–4 m tall. Leaves oblanceolate to oblong, blades 18–32 cm in length. Inflorescence 7–12 flowered, corolla white, sometimes tinged with pale purple. Occurs on shaded slopes and canyons in mesic forests at 530–730 m in the Wai‘anae Mountains, O‘ahu. Estimates suggest a population of several thousand individuals, with threats from feral pigs and goats and alien plant species.

4.165 

Cyanea pinnatifida

81

82

FLOWERING PLANTS

Shrubs 0.8–3.0 m tall, unbranched or sparingly branched. Leaves pinnately divided one-half to three-quarters of the distance to the midrib, blades 35–60 cm in length, upper surface glabrous, lower surface sparsely pubescent. Inflorescence 8–15 flowered, corolla greenish white with purplish stripes at the base. Rare in diverse mesic forest of the central Wai‘anae Mountains, O‘ahu.

Shrubs to 1.5 m tall, branched, stems smooth or with sharp prickles. Leaves smooth to sparsely prickly on both surfaces, broadly elliptic to obovate, blades 15–23 cm in length. Inflorescence 6–12 flowered, corolla white, sometimes with pale lilac stripes, often covered with small spines. Occurs in mesic forest at elevations of 460–850 m on West Maui.

Cyanea remyi Rock

hāhā, pua kala

hāhā

Cyanea solenocalyx Hillebr.

flowered, corolla, white with a longitudinal purple stripe on each lobe, barely emerging from the calyx. Rare in wet forest at elevations of 900–1,390 m on eastern Moloka‘i. Estimates suggest a population of less than 1,000 individuals, with threats from feral pigs, goats, deer, and alien plant species.

Cyanea spathulata (Hillebr.) A. Heller hāhā

ENDEMIC | SECURE

ENDEMIC | RARE

ENDEMIC | ENDANGERED

4.170  4.166 

Cyanea remyi

Shrubs 1–2 m tall, generally unbranched, stems smooth. Leaves broadly elliptic to oblong, blades 15–41 cm in length, leaf margins hardened and slightly toothed. Inflorescence 6–13 flowered, corolla dark purple. Occurs in wet forest at elvations of 360–933 m on Kaua‘i (Wailua, Wahiawa, and Wai‘oli valleys). A 2010 survey estimated no more than 24 individuals divided among 3 populations. Threats come primarily from feral pigs and alien plant species. Described in the Manual under the discussion of Cyanea truncata.

4.168 

Cyanea solenocalyx—habit

Cyanea spathulata

Branched shrub 2–3 m tall. Leaves narrowly oblanceolate or elliptic, 8–22 cm in length. Inflorescence 8–10 flowered, corolla white to greenish white, sometimes tinged with purple. Occurs in mesic to wet forest at 700–1,200 m elevation on western and south-central Kaua‘i (‘Ōpaewela, Kahōluamanu, Kōke‘e, Kalalau).

Cyanea st.-johnii (Hosaka) Lammers, Givnish & Sytsma hāhā

ENDEMIC | ENDANGERED

Cyanea scabra Hillebr. hāhā

ENDEMIC | SECURE

4.169 

4.167 

Cyanea scabra

Cyanea solenocalyx

Palmlike trees to 4 m tall with unbranched stems. The leaves are dimorphic with age: juveniles are prickly on both surfaces, while the adults are smooth above and unarmed below or only prickly along the midrib. Mature leaves oblong to obovate, 25–60 cm in length. Inflorescence 5–10

4.171 

Cyanea st.-johnii

Small woody shrubs, stems 30–60 cm in height. Leaves lanceolate to oblanceolate, blades 6–13 cm in length, lower leaf surface pubescent along the veins, margins strongly revolute. Inflorescence 5–20 flowered, corolla white with pale lilac on inner surfaces. Occurs in cloudswept wet forest

FLOWERING PLANTS

Last seen in the wild in 1983, but it has survived in cultivation, with some plants returned to exclosures in its native habitat to provide protection from feral pigs.

along the summit of the central Ko‘olau Mountains, O‘ahu. A 2007 report estimated 70 remaining individuals divided among 7 populations in the upper elevations of the Ko‘olau Mountains. Listed in the Manual as Rollandia st.-johnii.

Delissea—An endemic Hawaiian genus of

Cyanea superba (Cham.) A. Gray hāhā

ENDEMIC | ENDANGERED/EXTINCT

4.174 

4.172 

Cyanea superba subsp. superba

Palmlike trees 4–6 m tall. Leaves oblanceolate, blades 50–100 cm in length. Inflorescence 5–15 flowered, pendulous, densely pubescent, corolla white to cream with purple veins. Two subspecies have been described, with subsp. superba from mesic forest at elevations of 580–610 m in the northern Wai‘anae Mountains, O‘ahu. The last surviving individuals in the wild died in 2002, but the species survives today in cultivation. The subsp. regina formerly grew in the Ko‘olau Mountains and is thought to be extinct.

Cyanea tritomantha

Palmlike trees to 1.8–3.0 m tall. Leaves oblong to oblanceolate, prickly along the midrib beneath, blades 38–95 cm in length. Inflorescences 5–20 flowered, clustered among the leaves, corolla greenish white to black-purple outside but whitish to pale blue inside. An uncommon species in wet forest at elevations of 350–1,080 m in Waipi‘o Valley and on the windward slopes of Mauna Kea and Mauna Loa, Hawai‘i. Known from 16 populations totaling less than 400 plants. Threats come from feral pigs and cattle, ranching, and alien plant species.

Cyanea truncata (Rock) Rock hāhā

ENDEMIC | EXTINCT

Cyanea tritomantha A. Gray

15 species of shrubs or trees (one palmlike in growth form) with at least 6 and possibly 7 species extinct and the others rare and endangered. The name honors Jacques Delisse, a physician from Mauritius who served as a naturalist on an expedition to the South Pacific at the turn of the nineteenth century. These are plants of the dry to mesic forest. The flowers are curved, white or greenish, with several in an inflorescence. The corollas also have a dorsal knob. The seeds are large, white, and often wrinkled.

Delissea argutidentata (E. Wimm.) H. St. John ‘oha

ENDEMIC | ENDANGERED

Unbranched palmlike tree, 2–10 m tall. Stems woody, up to 5 cm diameter. Leaves variable in shape, 5–21 cm in length, arrayed in a dense apical rosette. Inflorescence 5–20 flowered, corolla greenish white. Once occurring at elevations of about 1,000 m in Acacia/Sophora woodland on the west and southwest slopes of Hualālai, Hawai‘i (see figure 8.1). Known today from a single individual. Listed in the Manual within the concept of Delissea undulata subsp. undulata.

Delissea kauaiensis (Lammers) Lammers ‘oha

‘akū, hāhā

ENDEMIC | ENDANGERED

ENDEMIC | VULNERABLE

4.175 

4.173 

Cyanea tritomantha—habit

Cyanea truncata

Shrubs 0.3–2.0 m tall, typically unbranched or with a few branches at the base, stems with small, sharp points. Leaves obovate, blades 22–60 cm in length, lower leaf surface pubescent with scattered small spines. Inflorescence 8–40 flowered, corolla white with highlights of magenta. Once occurred in mesic forest at elevations of 240–400 m on the windward slopes of the Ko‘olau Mountains, O‘ahu.

4.176 

Delissea kauaiensis

83

84

FLOWERING PLANTS

Unbranched palmlike tree, 2–10 m tall. Leaves ovate, blades 14–18 cm in length. Inflorescence 5–20 flowered, corolla greenish white. In mesic forest near Hanapēpē Falls, Kaua‘i. Thought to be extinct, it was rediscovered in 2002. Listed in the Manual as Delissea undulata subsp. kauaiensis.

Branched or unbranched shrubs, 1–3 m tall. Known today from a few small scattered populations in the Wai‘anae Mountains, O‘ahu. In 2008 there were 5 populations with about 40 individuals, with threats from feral goats and pigs. Listed in the Manual as Delissea subcordata var. kauaiensis.

Delissea rhytidosperma H. Mann

Lobelia—A large genus of about 300 spe-

‘oha

ENDEMIC | ENDANGERED

4.177 

Delissea rhytidosperma

cies of herbs, shrubs, and trees found in tropical and warm temperate regions around the world. The name honors Flemish botanist Mathias de l’Obel. In the Hawaiian Islands there are 15 endemic species that are shrubs. One group of closely related species has showy blue-violet flowers with tightly coiled petals. The other species have red to yellow or white flowers and are found in bogs or on the margins of bogs.

Lobelia gloria-montis Rock ENDEMIC | SECURE

4.181  Lobelia gloria-montis subsp. gloria-montis— habit

Lobelia dunbariae Rock ENDEMIC | RARE/EXTINCT

Branched shrubs 0.5–2.5 m tall. Stems are leafy in the upper portion with lanceolate to elliptic leaves blades 8–19 cm in length. Inflorescence 5–12 flowered, corolla greenish white with a prominent dorsal knob. Rare in diverse mesic forest at elevations of 300–1,000 m on Kaua‘i. Only 2 small populations are known.

Delissea waianaeensis Lammers ‘oha

ENDEMIC | ENDANGERED

4.182 

4.180 

4.178 

Delissea waianaeensis—habit

4.179 

Delissea waianaeensis

Lobelia dunbariae subsp. paniculata

Trailing shrubs with woody stems to 80 cm in length. Leaves linear to narrowly elliptic, 8–20 cm in length, sessile to subsessile. Inflorescence with 2–6 branches at its base, 15–25 cm in length, corolla blue. Occurs on cliff faces in wet forest at elevations of 1,000–1,200 m on Moloka‘i. Two subspecies are described, with subsp. paniculata with shorter calyx lobes pictured. There are 15 known localities with less than 2,000 individuals. Threats come from feral pigs, goats, deer, snails and slugs, and alien plant species.

Lobeila gloria-montis subsp. gloria-montis

Woody shrubs, stems to 5 m in length, unbranched with a dense apical rosette of oblanceolate to oblong leaves. Leaves oblanceolate to oblong, 8–24 cm in length, sessile. Inflorescence typically unbranched or rarely 3–7 branched from the base, 40–70 cm in length, corolla white, occasionally with purplish longitudinal stripes, rarely crimson. A striking species occurring in bogs and in wet forest at the margins of bogs at elevations of 1,350–1,750 m on West Maui and Moloka‘i. The subsp. gloria-montis occurs at Pāpa‘alā Pali on Moloka‘i and Pu‘ukukui on West Maui (see figure 2.1), while the subsp. longibracteata occurs on the rim of ‘Īao Valley and Haleakalā on Maui.

FLOWERING PLANTS

Lobelia grayana E. Wimm. ENDEMIC | SECURE

Woody shrub, stems unbranched to 2 m in length with a dense apical rosette of leaves. Leaves lanceolate to narrowly oblong, 12–30 cm in length. Inflorescence 2–5 branched from the base, 80–150 cm in length, corolla yellowish to greenish white with purple stripes. Occurs at elevations of 640–1,575 m in the Wahiawa and Wai‘ale‘ale bogs on Kaua‘i.

Lobelia niihauensis H. St. John ENDEMIC | ENDANGERED

Lobelia koolauensis (Hosaka & Fosberg) Lammers ENDEMIC | ENDANGERED

4.183 

Lobelia grayana

Woody shrub with erect stems 1–2 m in length and a dense apical rosette of leaves. Leaves linear, 10–24 cm in length, sessile to subsessile. Inflorescence unbranched, 10–40 cm in length, corolla blue. Occurs on cliffs in subalpine shrubland at 1,700– 2,300 m elevation in Haleakalā, East Maui.

4.186 

Lobelia niihauensis—habit

4.187 

Lobelia niihauensis

Lobelia kauaensis (A. Gray) A. Heller pu‘e

ENDEMIC | RARE

4.185 

4.184 

Lobelia kauaensis

Lobelia koolauensis

Woody shrub with a dense apical rosette of leaves, stems 30–100 cm in length and up to 4 cm in diameter. Leaves oblanceolate to oblong, 8–19 cm in length, sessile. Inflorescence unbranched or rarely 2–6 branched from the base, 40–70 cm in length, corolla crimson, rarely greenish or yellowish white. Occurs in cloudswept wet forest at elevations of 800–950 m along the crest of the northern Ko‘olau Mountains, O‘ahu. Only 2 populations are known with a total of about 280 individuals. Threats come from feral pigs and alien plant species. Listed in the Manual as Lobelia gaudichaudii subsp. koolauensis.

Woody shrubs with prostrate stems to 40 cm in length, branched at or below the middle and each branch with an apical rosette of leaves. Leaves narrowly elliptic, 7–15 cm in length, sessile to subsessile. Inflorescence unbranched, 12–15 cm in length, corolla magenta. Occurs on exposed mesic to dry cliff

85

86

FLOWERING PLANTS

faces at elevations of 125–725 m on Ni‘ihau (extinct), Kaua‘i, and the northern Wai‘anae Mountains, O‘ahu. Multiple populations remain on both Kaua‘i and O‘ahu with about 3,650 individuals reported, mostly on Kaua‘i. There are 14 populations and only about 400 individuals on O‘ahu.

The name is from the Greek trematos, hole, and the genus Lobelia. The fruit has perforations in the capsule, which upon drying allows the seeds to be shaken out in the wind, a unique adaptation in the family.

Trematolobelia macrostachys (Hook. & Arn.) Zahlbr. koli‘i

ENDEMIC | SECURE

Trematolobelia kaalae (O. Deg.) Lammers ENDEMIC | SECURE

Lobelia yuccoides Hillebr. pānaunau

ENDEMIC | RARE

4.191 

4.189 

4.188 

Lobelia yuccoides

Woody shrubs with erect stems and a dense apical rosette of leaves. Leaves linear, 24–35 cm in length, white-tomentose beneath, sessile to subsessile. Inflorescence unbranched, 6–100 cm in length, corolla blue to lilac. Occurs on dry ridges and canyons in diverse mesic forest at elevations of 750–1,200 m on Kaua‘i and in the Wai‘anae Mountains, O‘ahu.

Rollandia—Species formerly included in Rollandia are now placed in the genus Cyanea. Trematolobelia—An endemic Hawaiian

genus of 8 species of unbranched shrubs whose inflorescence comprises a terminal whorl of horizontal branches with the flowers arranged in one direction along the flowering axis. In 1 species, the flowers are borne in a single terminal branch.

Trematolobelia kaalae

Shrubs with an unbranched stem up to 2 m tall. Separated from the closely related T. macrostachys by the larger and more elongate shape of its floral tube and by a spring flowering phenology. Occurs in wet forests of the Wai‘anae Mountains and in particular on the upper slopes and summit of Mount Ka‘ala at elevations of 1,160–1,225 m. Listed in the Manual as T. macrostachys var. kaalae.

Trematolobelia macrostachys—dry fruits

Shrubs with unbranched stems up to 4 m tall. Leaves linear oblanceolate to narrowly elliptic, blades 9–24 cm in length. Inflorescences may have as many as 20 horizontal branches, corolla scarlet, rose, or pink. Flowers in summer and fall and is a favored nectar source for native honeycreepers. Occurs in wet forest and windswept ridges and cliffs at elevations of 550–1,200 m on O‘ahu (Ko‘olau Mountains), Moloka‘i, Lāna‘i, and both West and East Maui (see figure 2.1).

Trematolobelia singularis H. St. John ENDEMIC | ENDANGERED

Trematolobelia kauaiensis (Rock) Skottsb. koli‘i

ENDEMIC | SECURE

4.190 

Trematolobelia kauaiensis

Shrubs with an unbranched stem 1–3 m tall. Inflorescence composed of 5–10 horizontal branches, corolla scarlet. The plants die after flowering. Occurs in primarily wet forest at elevations of 650– 1,575 m on Kaua‘i.

4.192 

Trematolobelia singularis

FLOWERING PLANTS

Stems to 0.6–1.5 m tall. Leaves linear elliptic, blades 10–18 cm in length. Inflorescence unbranched, corolla violet. A rare plant at elevations of 900–915 m on the windswept summit of Konahuanui in the Ko‘olau Mountains, O‘ahu. As of 2009 there were 4 populations totaling about 360 individuals. Threats come from feral animals, alien plant species, slugs, and human disturbance.

Capparaceae (CAPER FAMILY)

A family of herbs, shrubs, and trees that produces mustard-oil glucosides and alkaloids. Key features are the presence of a gynophore (the stipe of the ovary is elongated within the flower) and usually irregular flowers often with many stamens. There are 19 genera and 510 species found in tropical and subtropical regions around the world. Economically of little importance except for 1 species cultivated for its edible flower buds. In the Hawaiian Islands represented today by the indigenous genus Capparis. The genus Cleome, once considered to be part of the Capparaceae, is now generally separated as the family Cleomaceae.

Capparis—A genus of climbing or sprawl-

ing shrubs or even small trees native to tropical and subtropical areas in the world. About 250 species are recognized. Capparis is the old Greek name for these plants. The caper of commerce is C. spinosa. In the Hawaiian Islands there is 1 endemic species that occurs in coastal areas.

Capparis sandwichiana DC. maiapilo, pua pilo, pilo

ENDEMIC | VULNERABLE

sunset. Scattered on coral, basaltic rocks, or in soil along the coast at elevations from sea level to 100 m on Midway and Pearl and Hermes atolls, Laysan (extinct), and all of the main islands.

Caryophyllaceae (PINK FAMILY)

A family of mostly annual or perennial herbs, rarely shrubs, or even small trees. Key features are opposite or whorled leaves with the leaf bases joined and sheathing the stem, and the fruit a single-chambered capsule that dehisces by valves or teeth. There are 85 genera and about 2,630 species found mostly in temperate and warm temperate areas of the world. The family is of economic importance because of the large number of ornamentals, such as carnations (Dianthus), baby’s breath (Gypsophila), and catchfly (Silene). In the Hawaiian Islands represented by 2 genera, Schiedea and Silene. At least 11 genera (including Spergula, Sagina, Drymaria, Polycarpon, and Arenaria) have become naturalized, with many invasive species.

4.195 

Schiedea adamantis

Erect shrubs to 80 cm tall. Leaves elliptic to oblanceolate, 1.5–3.0 cm in length. Flowers perfect or only female. Known from a single population on a dry, steep slope of Diamond Head Crater, O‘ahu, at an elevation of about 125 m. Less than 30 individuals survive, with threats from habitat loss.

Schiedea globosa H. Mann ENDEMIC | VULNERABLE

Alsinodendron—Four

species formerly placed in Alsinodendron have now been merged into the genus Schiedea.

Schiedea—An endemic Hawaiian genus of

34 species of shrubs, subshrubs, or vines, all representing single lineage. Many of the species are rare and endangered. The name commemorates a German physician, Christian Schiede. Four species having the trait of white fleshy sepals that enlarge in fruit were formerly segregated as the endemic genus Alsinodendron. Including Alsinodendron, the Manual described 26 species of Schiedea.

Schiedea adamantis H. St. John ENDEMIC | ENDANGERED

4.196 

4.193 

Capparis sandwichiana

Shrubs with decumbent or prostrate stems to 5 m long with fleshy, ovate, or elliptic leaves. The flowers, which have an irregular calyx and corolla, open after

4.194 

Schiedea adamantis—habit

Schiedea globosa

Low suffrutescent subshrubs, woody only at the base with prostrate to ascending stems to 20 cm in length. Leaves linear, thick, and somewhat fleshy, 3–10 cm in length. Plants typically dioecious. Occasionally a few perfect flowers occur. Occurs on steep, rocky slopes or cliffs in coastal habitats at elevations from sea level to 300 m on O‘ahu, Moloka‘i, Lāna‘i, Maui, and Hawai‘i.

87

88

FLOWERING PLANTS

Schiedea hookeri A. Gray ENDEMIC | ENDANGERED

4.197 

Schiedea hookeri

Trailing to weakly erect perennial herbs. Stems often in matted clumps, glabrous below, changing to small viscid hairs above. Leaves narrowly elliptic to lanceolate, 3–8 cm in length. Flowers perfect. Locally common in mesic forest at 350– 800 m elevation in the north and central Wai‘anae Mountains, O‘ahu, and Halea­ kalā on East Maui (extinct?). There are 17 known populations with less than 400 individuals. Threats come from feral goats and pigs and from alien plant species.

Schiedea kaalae Wawra ENDEMIC | ENDANGERED

Mountains (Makaleha, Pahole Gulch, Mokulē‘ia, Pu‘uhapapa, Huliwai) and Ko‘olau Mountains (Punalu‘u, Kaipapā‘u), O‘ahu. At last census there were only 9 small extant populations totaling less than 40 individuals. Threats have come from feral pigs and goats, alien plant species, and fire. Seedlings are reported to be consumed by slugs.

Erect to weakly erect shrub 30–60 cm tall. Leaves opposite, somewhat thickened, elliptic to linear, 3.3–4.5 cm in length. Plants dioecious. Scattered in occurrence in shaded dry to mesic forest, often on cliffs, at elevations of 640–730 m in the Wai‘anae Mountains, O‘ahu (Pālehua, Pu‘ukaua, Keawapilau Gulch, Mokulē‘ia).

Schiedea kealiae Caum & Hosaka

kuawāwaenohu

ENDEMIC | ENDANGERED

4.199 

Schiedea lychnoides Hillebr. ENDEMIC | ENDANGERED

Schiedea kealiae

Subshrubs, stems weakly ascending to sprawling to form loose clumps, 20–50 cm in length. Leaves opposite, thin, lanceolate to lanceolate-elliptic, conspicuously 3 nerved. Flowers unisexual and plants subdioecious. Scattered in remnant dry forest on steep slopes and ledges at elevations of 60–300 m in the northern Wai‘anae Mountains, O‘ahu.

Schiedea ligustrina Cham. & Schltdl. ENDEMIC | VULNERABLE

4.201 

Schiedea lychnoides

Weakly climbing or sprawling suffrutescent subshrubs, stems 40–300 cm in length, densely glandular. Leaves thin and membranous, ovate to elliptic, 3.5– 6.5 cm in length, 1.5–3.8 cm in width, densely glandular through out. Flowers perfect. A rare plant from the western margins of the Alaka‘i Swamp (Ke‘ākū Cave to Kahōluamanu), Kaua‘i, at elevations of 1,200–1,320 m. The most recent census found only 4 small populations with less than 50 mature plants. Threats come from feral pigs, alien plant species, and human disturbance. Listed in the Manual as Alsinodendron lychnoides.

Schiedea lydgatei Hillebr. ENDEMIC | ENDANGERED

4.198 

Schiedea kaalae

These nearly stemless plants arise from a woody caudex with the leaves tufted at the apex of the stem. Leaves thick and chartaceous, elliptic-lanceolate to nearly spathulate, 14–24 cm in length. Flowers perfect. Known from diverse mesic forests on steep slopes in shaded sites at elevations of 450–700 m in the Wai‘anae

4.202 

4.200 

Schiedea ligustrina

Schiedea lydgatei

Low, branched subshrubs with stems up to 40 cm in length. Leaves elliptic, 3

FLOWERING PLANTS

nerved, 2–6 cm in length. Flowers perfect. A rare species in mesic to dry forest at elevations of 600–650 m on Moloka‘i (Kamalō, Makakupa‘ia, and Popohua gulches, O‘oa). As many as 10,000 plants exist, but threats have come from fire, feral grazers, and alien plant species.

Schiedea mannii H. St. John ENDEMIC | VULNERABLE

Perennial herbs dying back to a woody base in dry seasons, stems ascending and sprawling, fleshy, unbranched, 50–100 cm in length. Leaves opposite, thin, broadly elliptic 13–20 cm in length. Flowers perfect, numerous in large open paniculate cymes. Rare in moist rocky sites in diverse mesic forest at elevations of 560–570 m on Kaua‘i (Mahanaloa and Ku‘ia valleys, Kalalau rim).

Schiedea menziesii Hook. ENDEMIC | VULNERABLE

Erect subshrub with branches 30–150 cm in length, stems white. Leaves opposite, green, sometimes tinged with purple, somewhat thickened, chartaceous, lanceolate to broadly or narrowly ovate. Flowers perfect. Once occurred in diverse mesic forest at elevations of 400–730 m in the Wai‘anae Mountains of O‘ahu (‘Ēkaha­ nui Gulch to Mokulē‘ia), Moloka‘i (eastern end, Kamalō, Kala‘e), and West Maui. Possibly extirpated now from Moloka‘i, West Maui, and the southern Wai‘anae Mountains. Currently there are 2 extant populations with about 40 mature individuals. Restoration efforts have added new plants to the population. Threats come from feral pigs and goats, alien plant species, military training, and fire.

Schiedea obovata (Sherff) W. L. Wagner & Weller ENDEMIC | ENDANGERED

4.205  4.203 

Schiedea mannii

Small shrubs, branched with stems 15–25 cm in length. Leaves opposite, somewhat thickened, linear to linear-elliptic, 2.5– 4.0 cm in length, 1 nerved. Flowers unisexual and plants dioecious. Scattered in open sites within diverse mesic forest at elevations of 400–800 m in the Wai‘anae Mountains (Pu‘uhapahapa, Pu‘ukawiwi, Mākua Valley, Mākua-Kea‘au Ridge, Ka­ maile‘unu Ridge).

Schiedea menziesii

Subshrubs with weakly ascending to sprawling stems forming loose clumps. Leaves thin and linear, 2–9 cm in length. Flowers perfect. Uncommon on ledges and cliffs in dry forest and shrubland at elevations of 30–340 m on Lāna‘i (Maunalei Valley) and West Maui (Mā‘alaea, Lahaina Luna Gulch).

Schiedea nuttallii Hook. ENDEMIC | ENDANGERED

Schiedea membranacea H. St. John ENDEMIC | ENDANGERED

4.207 

4.204 

Schiedea membranacea

4.206 

Schiedea nuttallii

Schiedea obovata

Erect, branched subshrubs to 1 m tall. Leaves thick and somewhat fleshy, elliptic, 4–11 cm in length. Inflorescence congested. Flowers perfect, adapted for bird pollination, petals absent, sepals enlarge and become fleshy at maturity, enclosing the fruiting capsule. A rare plant from slopes and ridges in diverse mesic forest at elevations of 550–800 m in the Wai‘anae Mountains, O‘ahu. Known today from only 2–3 populations with 14–44 individuals. Threats have come from alien plant species, habitat degradation from pigs, and fire. Listed in the Manual as Alsinodendron obovatum.

89

90

FLOWERING PLANTS

Schiedea salicaria Hillebr.

Schiedea trinervis (H. Mann) Pax & K. Hoffm.

ENDEMIC | ENDANGERED

ENDEMIC | ENDANGERED

in coastal soil pockets and cracks in cliff faces at elevations of 30–280 m on Nīhoa (West Palm Valley, Devil’s Slide). Extant populations are restricted to about 400 individual plants occurring in two rocky valleys.

Schiedea viscosa H. Mann ENDEMIC | ENDANGERED

4.208 

Schiedea salicaria

Erect subshrubs to 70 cm tall with stems forming loose clumps. Leaves narrowly elliptic to elliptic, 3–5 cm in length. Flowers perfect or all female. Scattered on ridges and steep slopes in remnant dry shrubland at elevations of 180–360 m on West Maui. Known from 3 populations with 500–1,000 individuals.

Schiedea spergulina A. Gray ENDEMIC | VULNERABLE

4.212 

4.210 

Schiedea trinervis

Erect, branched subshrubs to 80 cm tall. Leaves somewhat fleshy, elliptic-ovate, 6–12 cm in length. The flowers appear in an open compound inflorescence. Flowers without petals, 4 sepals enlarge, becoming fleshy and purple at maturity, enclosing the fruiting capsule. A very rare plant in wet forest at elevations of 900–1,230 m in the northern Wai‘anae Mountains, O‘ahu. Currently known from 2 populations with 192 individuals. Threats come from feral pigs and goats, and alien plant species. Listed in the Manual as Alsinodendron trinerve.

Schiedea verticillata F. Br. ENDEMIC | ENDANGERED

4.209 

Schiedea spergulina

Erect to weakly erect subshrubs to 60 cm tall. Leaves filiform, 3.0–6.5 cm in length. Flowers unisexual or perfect. Occurs on cliffs in diverse mesic forest at elevations of 180–250 m on Kaua‘i (Hanapēpē, Lāwa‘i Valley, Wahiawa, Olokele and Waimea canyons). The species is known from 7 populations and about 900 total individuals. It is threatened by habitat loss.

4.211 

Schiedea verticillata

Fleshy perennial herbs from an enlarged root. The fleshy leaves are in threes, are broadly ovate to elliptic, and are shed under dry summer conditions. Perfect flowers are the largest in the genus. Rare

Schiedea viscosa

Weakly climbing or sprawling subshrubs, stems to 3 m in length, densely covered by glandular hairs. Leaves thin and membranous, narrowly elliptic, 2.5–5.0 cm in length. Similar to A. lychnoides but differs in the width of the leaves and numbers of flowers per inflorescence. Occurs in diverse mesic to wet forests at elevations of 610–915 m on Kaua‘i (Waimea, Nāwaimaka, Wai‘alae, Kahōluamanu, west of Alaka‘i Swamp). Believed to be extinct until rediscovered in the early 1990s. Known from 4 fragmented populations with a total of less than 100 plants. Listed in the Manual as Alsinodendron viscosum.

Silene—A primarily north temperate

genus of about 700 species of mostly herbs and rarely subshrubs or shrubs, as in the Hawaiian species. The 5 petals are notably constricted in width at their base. The name probably honors Silenus, the foster father of Dionysus in Greek mythology. Many species are grown as ornamentals. In the Hawaiian Islands there are 7 endemic species of shrubs or subshrubs. These are thought to represent 2 distinct lineages, with S. hawaiiensis and S. struthioides forming one group and S. alexandri, S. cryptopetala, S. degeneri, S. perlmanii, and S. lanceolata the other. Two of these species, S. cryptopetala and S. perlmanii, are extinct in the wild. The xeromorphic shrub S. struthioloides is a relatively common alpine plant on Haleakalā, but the others are very rare or extinct.

FLOWERING PLANTS

Silene hawaiiensis Sherff ENDEMIC | RARE

in the Pohakuloa Training Area, Hawai‘i, with about 2,000 individuals. Currently there are only three populations totaling 100–130 individuals on O‘ahu. Threats come from fire, alien plant species, and feral grazing animals.

Silene perlmanii W. L. Wagner, D. R. Herbst & Sohmer ENDEMIC | EXTINCT

4.213 

Silene hawaiiensis

Sprawling suffrutescent shrubs, stems arising from a spindle-shaped root. Leaves slender and recurved, 6–15 mm in length. Scattered in occurrence in open dry areas on decomposed lava and ash at 900–1,300 m elevation or occasionally higher on Hawai‘i (Kīlauea, North Kona and Hāmākua districts, Saddle Road) (see figure 2.13). A recent census found 22 populations containing more than 8,000 individuals. Threats come from fire, alien plant species, and feral grazing animals.

4.217 

Silene lanceolata A. Gray catchfly

ENDEMIC | ENDANGERED

4.215 

Silene perlmanii

Subshrubs, usually many branched from the base, often forming clumps. An extremely rare species known from a single population on a cliff face in diverse mesic forest at an elevation of 790 m between Palikea and Pōhākea Pass in the Wai‘anae Mountains, O‘ahu. Now considered to be extinct, but there are outplantings of individuals raised in cultivation.

Silene struthioloides A. Gray 4.214 

Silene lanceolata

Subshrubs with erect to ascending stems to 50 cm in length and oblanceolate to lanceolate leaves. One of 5 white-flowered species native to the Islands. Rare in dry areas and mesic shrubland at elevations of 330–1,900 m in the southern Wai‘anae Mountains of O‘ahu, Moloka‘i (Pu‘ukolekole), and Hawai‘i (Pu‘uahi, Mauna Kea) and previously from Kaua‘i and Lāna‘i. The majority of the known plants occur

ENDEMIC | SECURE

4.216 

Silene struthioloides—habit

Silene struthioloides

Many-branched shrubs with stems from a taproot and awl-shaped, rigid leaves. The flowers open in the early evening, greenish when fresh but fading to maroon on drying. Occurs in subalpine to alpine shrubland at elevations of 2,150–3,500 m on Haleakalā, East Maui, and on Mauna Kea, Hawai‘i, with a single collection from Hualālai.

Celastraceae (BITTERSWEET FAMILY)—The genus Perrotettia, once con-

sidered to be a member of the Celastraceae, has now been placed in the Dipentodontaceae.

Chenopodiaceae (GOOSEFOOT FAM-

ILY)—The Chenopodia­ceae has now been merged into the Amaranthaceae.

Cleomaceae (SPIDER PLANT)—The members of the Cleomaceae were formerly placed in the Capparaceae, but recent molecular data suggest their segregation as a separate family or alternatively as part of a broader concept of the Brassicaceae. Cleome spinosa has sometimes been considered as indigenous to the Hawaiian Islands but is questionably native. It has not been collected since 1865 and is believed to be extinct in the Islands.

91

92

FLOWERING PLANTS

Convolvulaceae

(MORNING-GLORY FAMILY)

Erect or twining herbs, vines, lianas, shrubs, or small trees. Some key features are the presence of milky sap, plants often with large rootstocks, and a funnel-shaped corolla of fused petals; flowers typically perfect. There are 52 genera and about 1,650 species that are found in tropical and subtropical regions around the world. Cuscuta (dodder) is now included in this family. Economically the family is important because of the sweet potato and the large number of ornamentals in the genera Ipomoea, Convolvulus, Merremia, and Porana. Represented in the Hawaiian Islands by the indigenous genera Bonamia, Cressa, Ipomoea, and Jacquemontia.

total population is thought to be only a few hundred individuals. It has declined in number from habitat loss due to development and agriculture and from impacts of fire, feral grazing animals, and alien plant species. On O‘ahu the species has declined to 12–13 populations and fewer than 60 individuals.

Cressa—A small genus of 4 species of halophytic herbs or subshrubs widespread in the tropics and temperate areas. The name is from the Greek, kris, meaning from Crete, since the type species was described from Crete. In the Hawaiian Islands, the single indigenous species present is less common compared to other parts of its geographic range.

Cressa truxillensis Kunth INDIGENOUS | SECURE

Bonamia—A pantropical genus of 45 spe-

cies of lianas and occasionally vines or subshrubs. The name honors Francois Bonami, a French botanist and physician of the eighteenth century. In the Hawaiian Islands there is a single endemic species, which is considered rare. It is an unusual member of the genus, once having been described as an endemic genus Perispermum.

Bonamia menziesii A. Gray ENDEMIC | ENDANGERED

4.219 

Cressa truxillensis

Subshrubs or slightly woody herbs with erect to decumbent stems and fleshy leaves. The flowers are usually clustered at the tips of the branches. Widespread in the southwestern United States, Mexico, South America, and in the Hawaiian Islands, where it occurs on drying mudflats and sand along beaches on O‘ahu, Moloka‘i, and Kaho‘olawe.

4.218 

Bonamia menziesii

Lianas with twining stems up to 10 m long. Leaves soft, coriaceous, covered with yellowish brown hairs. Flowers usually solitary, covered with brownish hairs. A rare plant in dry to mesic forest at elevations of 150–625 m on all of the main islands except Ni‘ihau and Kaho‘olawe, but extinct on Moloka‘i. Although still present on most of the main islands, the

Cuscuta—A genus of 45 species of achlorophyllous parasitic herbs. The short-lived twining branches are a distinct orange color and attached to the host by haustoria. The genus includes about 150 species with a cosmopolitan distribution but with a center of diversity in the New World. Many species are considered to be agricultural pests. The name is derived from the Arabic work kuskut, meaning a tangle of hair. Represented in the Hawaiian Islands by 1 native and 1 naturalized species.

Cuscuta sandwichiana Choisy

kauna‘oa, kauna‘oa kahakai, dodder ENDEMIC | SECURE

4.220 

Cuscuta sandwichiana

Parasitic twining vines with scalelike reduced leaves, yellow to yellowish orange. Occurs on a wide variety of host plants that grow along the coastal areas. On all of the main islands except Kaho‘olawe.

Ipomoea—A large genus of vines, shrubs, or trees that are found throughout the tropics and subtropics of the world. About 650 species have been described, with 1 endemic and 4 indigenous species occurring in the Hawaiian Islands. The name is from the Greek ips, worm, and homoios, similar to, because the twining habit of many species is wormlike. Ipomaea cairica was introduced by the Polynesians and is discussed in chapter 6. Apart from the sweet potato, several members of this genus are grown as ornamentals. The Prince Kūhiō vine, Ipomoea horsfallii, is commonly planted in the Hawaiian Islands. Both Ipomoea alba and I. violacea are widely naturalized on the main islands.

Ipomoea imperati (Vahl.) Griseb. hunakai, morning glory INDIGENOUS | SECURE

4.221 

Ipomaea imperati

Perennial vines with trailing stems that root at the nodes, up to 4 m or more in length. Leaves variable in shape from linear to ovate, 1.5–4.0 cm in length, glabrous, somewhat fleshy. Pantropical in distribution. Occurs on beaches and

FLOWERING PLANTS

sand dunes from sea level to elevations of 20 m on Ni‘ihau, Kaua‘i, O‘ahu, Moloka‘i, and Maui.

Ipomoea indica (J. Burm.) Merr.

Ipomoea pes-caprae (L.) R. Br. subsp. brasiliensis (L.) Ooststr.

pōhuehue, puhuehue, beach morning glory INDIGENOUS | SECURE

koali ‘awa, koali, morning glory

sides of all of the main islands. The roots and leaves were used by the Hawaiians as a cathartic and also to treat thrush in infants. Listed in the Manual as Jaquemontia ovalifolia.

Cucurbitaceae

INDIGENOUS | SECURE

(GOURD FAMILY)

4.224 

4.222 

Ipomoea indica

Vines with twining stems, somewhat woody at the base, up to 5 m long. Leaves broadly ovate, entire to 3 lobed, 5–9 cm in length, densely pubescent, especially on the lower surface. Flowers usually blue or purple, turning pink in the afternoon, rarely white. Common in dry disturbed areas at elevations from sea level to 150 m on Kure and Midway atolls, Lisianski, Laysan, Nīhoa, and on all of the main islands. The Hawaiians utilized the leaves and roots in plasters and poultices for wounds and also in treating broken bones. The seeds were used as a cathartic.

Ipomaea littoralis (L.) Blume INDIGENOUS? | SECURE

Ipomoea pes-caprae

Pantropical vines with long trailing stems often rooting at the nodes. Leaves fleshy, often arranged on one side of the stem, and variable in shape but usually with an apical notch, 5–11 cm in length, glabrous. Flowers pink to purple, rarely white. Occurs commonly on beaches and occasionally inland on Kure and Midway atolls, Lisianski, Laysan, French Frigate Shoals, Nīhoa, and all of the main islands. The roots and leaves of this plant were said to be used as famine food.

Jacquemontia—A genus of 120 species of vines, herbs, or subshrubs in the tropics and subtropics, especially the Americas. The name honors Victor Jacquemont, a nineteenth-century French botanical explorer. In the Hawaiian Islands, the endemic species was once considered to be a subspecies of the widespread Jaquemontia ovalifolia.

Jacquemontia sandwicensis A. Gray kākuaohi‘iaka, kaupo‘o, pā‘ūohi‘iaka ENDEMIC | SECURE

Climbing or trailing annual or perennial herbs, rarely somewhat woody or arborescent. Key features are the typically viny habit, the presence of tendrils at the leaf nodes in most genera, and monoecious (rarely bisexual) with an inferior ovary. There are 121 genera and about 760 species found throughout the tropics and also in temperate regions around the world. Economically, the family is the source of melons, pumpkins, squash, cucumbers, and gherkins. A few are grown as ornamentals, and the bottle gourd (Lagenaria siceraria) is widely cultivated in the tropics for its hard-shelled fruit. In the Hawaiian Islands, Sicyos is the only indigenous genus, but the balsam pear (Mormodica), the teasel gourd (Cucumis), and the ivy scarlet gourd (Coccinia) are naturalized, with the latter a highly invasive species.

Sicyos—A genus of about 25 species of annual or perennial climbing or trailing herbs, often woody at the base, and distributed in North America, the southwestern Pacific, New Zealand, Australia, and the Hawaiian Islands. Flowers monoecious. There are 14 endemic Hawaiian species, which probably originated from a single South American lineage. Sicyos is the Greek word for cucumber. The fruit is variously pubescent, spiny, bristly, or ribbed. The plants have no economic importance.

Sicyos anunu (H. St. John) I. Telford ‘ānunu

ENDEMIC | SECURE

4.223 

Ipomoea littoralis

Vines, stems typically prostrate, rooting at the nodes, woody at the base. Leaves thin, coriaceous, ovate, 1–10 cm in length. Flowers usually solitary, corolla lavender to pinkish purple. Occurs in coastal sites on O‘ahu, Moloka‘i, Maui, and once reported from Hawai‘i. Widely distributed across the Pacific and Indian oceans and questionably indigenous in Hawai‘i.

4.225 

Jacquemontia sandwicensis

Vines with prostrate stems often rooting at the nodes, up to 3 m long. Leaves thick and sometimes fleshy, elliptic to suborbicular, 1–6 cm in length. Stems and leaves with branched hairs. Flowers pale blue to white. A common plant in coastal habitats, particularly the leeward

4.226 

Sicyos anunu

93

94

FLOWERING PLANTS

Climbing annual herbs with stems to 8 m long. Leaves broadly ovate, shallowly to deeply palmately lobed, 7–16 cm in length. Pistillate flowers 35–65 per head. Fruit fusiform with a long beak. Occurs in dry forest or subalpine forest at elevations of 350–1,940 m on Lāna‘i and Hawai‘i.

Sicyos lasiocephalus Skottsb.

Sicyos pachycarpus Hook. & Arn.

ENDEMIC | SECURE

ENDEMIC | SECURE

‘ānunu

‘ānunu, kūpala

Sicyos herbstii (H. St. John) I. Telford ‘ānunu

ENDEMIC | SECURE

4.231 

4.229 

4.227 

Sicyos herbstii

Climbing annual herbs with stems to several m long. Leaves broadly ovate, shallowly palmately 5–7 lobed, 3.5–7.0 cm in length. Pistillate flowers 6–9 per head. Fruit ovoid, 5 ridged, covered with multicellular hairs. Occurs in shrub communities at elevations from sea level to 500 m on the leeward side of Kaua‘i (Makaweli to Barking Sands and Polihale). A single collection from Moloka‘i questionably represents this species.

Sicyos lasiocephalus

Climbing annual vines with stems up to several meters in length, densely covered with multicellular hairs. Leaves suborbiculate to broadly cordate, 5 angled to shallowly to deeply 5 lobed, 3.5–9.0 cm in length. Pistillate flowers 30–50 in a dense head. Fruit ellipsoid, densely covered with silky hairs. Typically occurs in open ‘ōhi‘a forest at elevations of 600–800 m on Moloka‘i (Kalama‘ula) and Hawai‘i (north slope of Hualālai, leeward slope of the Kohala Mountains).

Sicyos maximowiczii Cogn. puaokama

Sicyos pachycarpus

Climbing vines, possibly with perennial rootstocks, stems up to several m long. Leaves broadly ovate-cordate to kidney shaped, shallowly 5–7 lobed, 7–26 cm in length. Pistillate flowers 10–50 per head. Fruit ovoid, black or brown, abruptly contracted into a short beak. An extremely variable species that is widespread in shrubland coastal communities, dry forest, and in introduced alien vegetation on the lower leeward slopes of all of the main islands at elevations from sea level to 900 m. It has also been collected from Laysan and Nīhoa.

Sicyos waimanaloensis H. St. John ‘ānunu

ENDEMIC | RARE

ENDEMIC | SECURE

Sicyos hispidus Hillebr. ‘ānunu

ENDEMIC | SECURE

4.232 

4.228 

Sicyos hispidus

Climbing annual vines with stems up to 6 m long covered with multicellular glandular hairs. Leaves broadly ovate-cordate, usually 5 angled or shallowly palmately 5 lobed, 7–16 cm in length. Pistillate flowers 6–12 per head. Fruit ovoid, densely pubescent with pale multicellular glandular hairs. Occurs in dry forest or introduced vegetation at elevations from sea level up to 800 m on Moloka‘i, Lāna‘i, Maui, and Hawai‘i (North Kona).

4.230 

Sicyos maximowiczii

Trailing or clambering annual vine with stems up to 10 m in length. Leaves kidney shaped to broadly ovate cordate, 5–7 angled to shallowly lobed, 6–12 cm in length. Pistillate flowers 10–25 per head. Fruit obovoid, densely covered with silky glandular hairs. Occurs across the Northwestern Hawaiian Islands on coral sands near sea level. Known from Kure Atoll, Pearl and Hermes Atoll, Lisianski, Laysan, and Ni‘ihau and once present on O‘ahu.

Sicyos waimanaloensis

Climbing annual herbs with stems to several m long, glabrous. Leaves broadly ovate-cordate or kidney shaped, 5–7 lobed, 4–14 cm in length. Pistillate flowers 7–15 per head. Fruit ovoid, lance shaped, tapering into a long beak. Rare in open sites or shrubland communities at elevations of 50–650 m on Kaua‘i (Waimea Canyon), O‘ahu (now extinct), and Moloka‘i.

Cuscutaceae

(DODDER FAMILY)—

The Cuscutaceae, including the Hawaiian genus Cuscuta, has been merged into the Convolvulaceae.

FLOWERING PLANTS

Dipentodontaceae (DIPENTODON FAMILY)

Trees or shrubs, with semievergreen or deciduous leaves; flowers perfect or dioecious. Two genera and about 16 species in tropical Asia to Mexico, Peru, Pacific Islands, and extending into warm temperate East Asia. Represented in the Hawaiian Islands by the indigenous genus Perrottetia.

Perrottetia—A genus of shrubs or small

trees with 15 species distributed in tropical America, Queensland, China, Indonesia, Malaysia, New Guinea, to the Pacific Islands. Leaves are alternate with a prominent midrib. Named in honor of George Perrottet, a French botanist and gardener. One species occurs in the Hawaiian Islands. This genus was formerly placed in the Celastraceae.

Perrottetia sandwicensis A. Gray olomea, pua‘a olomea

aquatic (Aldovandra). Key features are the insectivorous nature of the leaves and the bisexual, primarily 5-merous flowers. There are 3 genera and about 105 species. A few species of sundew (Drosera) and the Venus flytrap (Dionaea) are occasionally cultivated as novelties. In the Hawaiian Islands, only 1 indigenous species of the genus Drosera occurs in bogs.

Drosera—A genus of about 100 species of

cosmopolitan herbs with many occurring in the Southern Hemisphere, particularly Australia. The name is from the Greek droseros, which means dewy and refers to the appearance of the glandular hairs on the leaves. In the Hawaiian Islands, 1 species occurs on Kaua‘i in bogs, probably brought in on the feet of migratory golden plovers at some point in the not too distant past.

and date-plums—and as timber because of the very dense wood. In the Hawaiian Islands represented by the indigenous genus Diospyros.

Diospyros—A pantropical genus of about

550 species of trees and shrubs. The Hawaiian species are thought to have a Malesian origin. The name comes from the Greek dios, Zeus, and pyros, grain, in reference to the edible fruit of some species. In the Hawaiian Islands, there are 2 endemic species and several cultivated species.

Diospyros hillebrandii (Seem.) Fosberg lama, ēlama, ebony, persimmon ENDEMIC | SECURE

Drosera anglica Huds. mikinalo, sundew

INDIGENOUS | SECURE

ENDEMIC | SECURE

4.235 

4.233 

Perrottetia sandwicensis

Shrubs to small trees 2–6 m tall, branches red to green, short and stiff, often drooping at higher elevations. Leaves ovate, elliptic or oblong-elliptic, 7–19 cm in length, with reddish veins and petioles. Polygamodioecious, congested inflorescences bear many tiny flowers, followed by small red fruit. A common understory tree in wet forest at elevations of 300–1,250 or higher on all of the main islands except Ni‘ihau and Kaho‘olawe. The Native Hawaiians made fire by rubbing pieces of the wood against the soft wood of the hau, Hibiscus tiliaceus.

Droseraceae

(SUNDEW FAMILY)

Annual or perennial herbs or subshrubs that are insectivorous, using glandular tentacle hairs that secrete mucilage to trap insects. One member is a submerged

4.234 

Drosera anglica

Small stemless perennial herbs arising from short rootstocks. Leaves covered with sticky, glandular hairs that trap small insects. Occurs in bogs on Kaua‘i but curiously absent from bogs on Moloka‘i, Maui, and Hawai‘i. Elsewhere it is native to bogs in the Northern Hemisphere.

Diospyros hillebrandii

Trees 4–7 m tall or more. Leaves coriaceous, oblong-elliptic, 8–16 cm in length, dark green, with upper surface conspicuously pitted, both leaf surfaces glabrous. Fruit pale orange in color. Scattered to common in species-rich mesic forest at elevations of 150–760 m on Kaua‘i and O‘ahu.

Diospyros sandwicensis (A. DC.) Fosberg lama, ēlama, ebony, persimmon ENDEMIC | SECURE

Ebenaceae

(EBONY FAMILY)

A family of evergreen and deciduous woody shrubs and trees, widely distributed in tropical to warm temperate regions, with 3 genera and about 575 species. Key family features are the presence of red, green, or black heartwood, the latter the source of ebony. Plants dioecious with fleshy to leathery berries on the females. Economically the family is important for the fruits—persimmons

4.236 

Diospyros sandwicensis

Trees to 10 m tall, often with galls on the stems. Leaves coriaceous, thick, pale green with inconspicuous fine reticulate pits, and generally arranged all in one plane, lanceolate to falcate, 3–9 cm in length, lower leaf surface with stiff hairs. Ellipsoid

95

96

FLOWERING PLANTS

fruit is reddish orange to yellow. A variable plant throughout its range in the Hawaiian Islands. Occurs in dry to mesic forest, where it is sometimes dominant at elevations from 5 to 1,220 m. On all of the main islands except Ni‘ihau and Kaho‘olawe. The wood of the plant was used medicinally and to fence off sacred areas.

Elaeocarpaceae

(ELAEOCARPUS FAMILY)

Trees and shrubs from tropical and subtropical areas except absent from continental Africa. Key features that differentiate this family from the closely related Malvaceae (Tiliaceae) are the pubescence of simple or glandular hairs and the petals, when present, not being imbricate or overlapping. There are 12 genera and about 600 species. A few representatives of Elaeocarpus, Crinodendron, Aristotelia, and Muntingia are occasionally cultivated. In the Hawaiian Islands represented by the indigenous genus Elaeocarpus. The calabura (Muntingia calabura) is occasionally cultivated in the Islands.

Elaeocarpus—A genus of about 350 species

of trees or shrubs distributed from Madagascar and Mauritius to eastern Asia, Malesia, New Zealand, Australia, and the Pacific, including 1 species in the Hawaiian Islands. Some are cultivated as ornamentals, one yields a blue-black dye, another is the source of karanda nuts in Australia, and the fruit of E. serratus is used as a condiment in curries. The name is from the Greek elaia, olive, and karpos, fruit, because the fruit closely resembles olives. In the Hawaiian Islands, there is 1 endemic species confined to O‘ahu and Kaua‘i.

Elaeocarpus bifidus Hook. & Arn. kalia

ENDEMIC | SECURE

Trees to 10 m tall, apex of young branches viscid. Leaves ovate to lanceolate, 11–18 cm in length, margins crenate, pale green to pale orange petioles. Flowers 5–8 in racemes 4.5–6.0 cm in length, inflorescences are often attacked by mites causing a red “witches broom.” Occurs in diverse mesic to wet forest and the margins of bogs at elevations of 900– 1,220 m on Kaua‘i and O‘ahu. The bark was formerly harvested to provide cordage, and the wood was used for house frames.

Epacridaceae

Elaeocarpus bifidus

ENDEMIC | SECURE

(EPACRIS FAMILY)—

The former Epacridaceae, which included the Hawaiian genus Leptecophylla, has now been merged into the Ericaceae.

Ericaceae

(HEATH FAMILY)

Terrestrial or epiphytic shrubs, lianas, and small trees. Key features are the often urn-shaped corollas, the stamens arising from a nectar-bearing disc, and an often 4-parted inferior ovary. There are 117 genera and about 3,850 species in temperate and subtropical areas and in mountains in the tropics. A large group of Australian and Southern Hemisphere genera was formerly separated into a distinct family as the Epacridaceae. Economically the family is important for fruits such as blueberries and cranberries (Vaccinium), for many cultivated ornamentals such as azaleas and rhododendrons (Rhododendron), heaths (Erica), heathers (Calluna), mountain laurels (Kalmia), and andromedas (Pieris) and as foliage plants in the florists trade, specifically salal leaves (Gaultheria shallon). The indigenous genera Leptecophylla and Vaccinium occur in the Hawaiian Islands.

Leptecophylla—A genus of 13 species of

4.237 

Leptecophylla tameiameiae (Cham. & Schltdl.) C. M. Weiller pūkiawe

erect shrubs to small trees found primarily in Australia and Tasmania, with a few distributed in southeastern Asia, New Zealand, New Caledonia, and the Hawaiian Islands. The name is from the Greek lepto, small or fine, and phyll, leaf. In the Hawaiian Islands, there is 1 highly variable species that occurs in a wide variety of habitats. A high-elevation species with a compact habit was formerly recognized as distinct.

4.238 

Leptecophylla tameiameiae

Compact to slender, erect and spreading shrubs (trailing in bogs), 1–2 m in height or rarely arboreal up to 5 m tall. The closely spaced, coriaceous leaves are variable in shape and size but typically 6–13 mm in length and 3 mm wide, with a sharp, acute tip. Tiny flowers are pink or white. Fruit somewhat fleshy berries, dark red to pink or white. A common and morphologically plastic species often becoming dominant in mesic forest, montane wet forest, alpine shrubland and bogs at a broad range of elevations from 330–2,320 m on all of the main islands except Ni‘ihau and Kaho‘olawe but possibly on these islands in the past. The berries of the plant are often used in lei making. This species was listed in the Manual as Styphelia tameiameiae.

Vaccinium—A genus of at least 450 species of terrestrial (sometimes epiphytic) shrubs, small trees, or lianas distributed in north temperate areas or subtropical and tropical mountains throughout the world. The name is derived from the Latin vaccinus, of cows, referring to an old Germanic folk name. Cranberries, huckleberries, bearberries, and blueberries are edible fruits of species in this genus. A few have ornamental value. In the Hawaiian Islands represented by 3 endemic species, one of which is highly morphologically variable.

FLOWERING PLANTS

Vaccinium calycinum Sm.

Vaccinium reticulatum Sm.

ENDEMIC | SECURE

ENDEMIC | SECURE

‘ōhelo kau lā‘au, ‘ōhelo

4.239 

‘ōhelo

Vaccinium calycinum

Erect shrubs to 5 m tall. The ovate to obovate leaves are deciduous. Flowers are green, yellowish green, or reddish green. Common in wet forest and bogs at elevations of 500–1,800 m on all of the main islands except Ni‘ihau and Kaho‘olawe. The bright red berries are bitter and are rarely used in the making of jams and jellies.

Vaccinium reticulatum—Maui; this form was formerly recognized as V. berberidifolium

4.241 

Vaccinium dentatum Sm. ‘ōhelo

ENDEMIC | SECURE

4.240 

4.242 

Vaccinium reticulatum—Mauna Kea

4.243 

Vaccinium reticulatum—Kīlauea

Vaccinium dentatum

Decumbent or sprawling shrubs up to 2 m tall. The coriaceous elliptic or narrowly elliptic leaves are persistent. Flowers are red with greenish lobes, green with red stripes, or pink with greenish lobes. The bright red fruit is widely used in the making of jams and jellies. Occurs along the edges of wet forest, bogs, swamps, or exposed ridges at elevations of 700– 1,200 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Variable shrubs with coriaceous, persistent, ovate to obovate leaves with leaf margins ranging from entire to strongly toothed. The flowers are red, yellow, yellow with red stripes, or greenish yellow. The edible fruits may be red, black, dull yellow, yellow-green, pink, or sometimes purplish. A common shrub of disturbed sites at elevations of 640–3,700 m on Maui and Hawai‘i and rare on Kaua‘i, O‘ahu, and Moloka‘i. Frequent on lava flows and cinder beds in the Kīlauea area. Morphologically variable. Plants formerly named as Vaccinium berberidifolium are now considered to be part of this complex. The berries are used in the making of jams and jellies. ‘Ōhelo berries were sacred to Pele. Branches of the plant with berries were thrown into the crater of Halema‘uma‘u before consumption so as not to anger the goddess.

Euphorbiaceae (SPURGE FAMILY)

Annual or perennial herbs, shrubs, stem succulents, trees, and sometimes even vines. Cosmopolitan in distribution including both temperate and tropical regions but absent from the cold Arctic. Key features are the presence of milky sap (in many genera), unisexual flowers borne on the same or separate plants, and the usually 3-parted superior ovary. There are 229 genera and about 6,500 species. Economically the family is important for the production of rubber (Hevea), tung oil (Aleurites), castor oil (Ricinus), cassava, and tapioca (Manihot) and for ornamentals in the genus Euphorbia and Codiaeum. In the Hawaiian Islands represented by the indigenous genera Claoxylon and Euphorbia. The segregate genus Chamaesyce was formerly used for 16 endemic species, but these have all now been placed back into Euphorbia. This clade is sister to a group of annuals from Texas and northern Mexico. The genera Antidesma, Flueggea, Phyllanthus and related genera, once considered as members of the Euphorbiaceae, are now placed within the Phyllanthaceae. Introduced genera that have become naturalized include Aleurites (kukui), Croton, Jatropha, Mallotus, Macaranga, Manihot, Homalanthus, and Ricinus.

Chamaesyce—The 16 Hawaiian species of Chamaesyce, along with all others

97

98

FLOWERING PLANTS

formerly included in this genus, have now been placed within Euphorbia.

Claoxylon—A paleotropical genus of about

75 species of shrubs and trees. The name is from the Greek clao, break, and xylon, wood, in reference to the fact that the plants are soft wooded. In the Hawaiian Islands represented by 1 somewhat variable species. The male flowers are arranged in clusters, while the females are solitary.

species occur in a wide variety of habitats, from coastal to dry, mesic, and wet forest and even bogs. This is an excellent group to illustrate the concept of adaptive radiation, as the Hawaiian species are unusual in being woody plants utilizing C 4 metabolism.

Euphorbia atrococca A. Heller

‘akoko, ‘ekoko, kōkōmālei, spurge ENDEMIC | SECURE

4.247 

Claoxylon sandwicense Mull. Arg. po‘olā

Shrubs or small trees to 8 m tall. Leaves variable in shape and distichous. Eight varieties are recognized, with var. amplectens widespread on the main islands and the var. kaenana restricted to O‘ahu. This is the most variable and widespread species in the Hawaiian Islands, occurring in coastal habitats and in mesic forest at elevations from sea level to 1,800 m on all of the main islands and Nīhoa. Listed in the Manual as Chamaesyce celastroides.

ENDEMIC | SECURE

4.244 

Euphorbia clusiifolia Hook. & Arn.

Claoxylon sandwicense

Soft-wooded monoecious shrubs or trees to 5 m tall, with oblong, oblanceolate to elliptic leaves. In mesic to sometimes dry forest at elevations of 550–650 m or rarely higher on all of the main islands except Ni‘ihau and Kaho‘olawe.

Euphorbia—A large genus of about 2,400 species of annual or perennial herbs, shrubs, trees, and sometimes stem succulents (notably in Africa); found throughout the world but predominantly in tropical, subtropical, or warm temperate areas. The name reputedly honors Euphorbus, a Mauretanian physician in the first century AD. The latex of most of the species is toxic and can cause skin rashes in sensitive individuals. Some of the succulent species are popular as ornamentals. The poinsettia, a common Christmas ornamental, is widely cultivated in tropical regions. In the Hawaiian Islands there are 17 endemic species, with all but one of them (E. haeleeleana) formerly placed in Chamaesyce, a previously recognized cosmopolitan genus of about 250 species of herbs, shrubs, or small trees that are predominantly New World. The Hawaiian Euphorbia (Chamaesyce)

Euphorbia celastroides var. kaenana—habit

‘akoko, ‘ekoko, koko, spurge ENDEMIC | SECURE

4.245 

Euphorbia atrococca

Shrubs or small trees up to 3 m with leaves that are somewhat variable in shape and decussate. Flowers clustered at the ends of the branches or solitary in the leaf axils. Rare in dry forest to diverse mesic forest at elevations of 400–900 m on western Kaua‘i. Listed in the Manual as Chamaesyce atrococca.

Euphorbia celastroides Boiss.

‘akoko, ‘ekoko, kōkōmālei, spurge

ENDEMIC | SECURE/VULNERABLE/ENDANGERED/ EXTINCT

4.248 

4.246 

Euphorbia celastroides var. amplectens

Euphorbia clusiifolia

Shrubs with erect to scandent stems up to 3 m long. Leaves oblanceolate to oblong, usually decussate. Occurs in mesic to wet forest at elevations of 300–600 m on the western slopes of the Ko‘olau Mountains, O‘ahu. Listed in the Manual as Chamaesyce cluisiifolia.

FLOWERING PLANTS

Euphorbia degeneri Sherff

Euphorbia haeleeleana D. R. Herbst

ENDEMIC | SECURE

ENDEMIC | ENDANGERED

‘akoko, ‘ekoko, koko, spurge

spurge

Shrubs with scandent stems to 4 m long. The elliptic to oblanceolate leaves are decussate. Flowers are usually in dense, branched clusters. Occasionally in diverse mesic or wet forest, at elevations of 400–1,100 m on northwestern Kaua‘i. The most recent census reported 4 extant populations with less than 400 plants. Threats come primarily from feral grazing animals. Listed in the Manual as Chamaesyce halemanui.

Euphorbia herbstii (W. L. Wagner) Oudejans ‘akoko, ‘ekoko, koko, spurge ENDEMIC | ENDANGERED

4.251  4.249 

Euphorbia degeneri

Subshrubs with prostrate to decumbent stems, often tortuous and gnarled. Leaves orbicular to elliptic, distichous. Flowers at the ends of the branches. Scattered in coastal and strand vegetation from sea level to 50 m elevation on all of the main islands except Lāna‘i and Kaho‘olawe. Listed in the Manual as Chamaesyce degeneri.

Euphorbia deppeana Boiss. ‘ekoko, koko, spurge

ENDEMIC | ENDANGERED

Euphorbia haeleeleana

Small trees 3–14 m tall with elliptic leaves. Plants dioecious. Although locally common in diverse mesic forest at elevations of 350–650 m on Kaua‘i, it was not discovered until the late 1960s. It also occurs in two pockets of remnant dry forest in the Wai‘anae Mountains, O‘ahu. Populations have been declining and there are currently 6 populations with an estimated 65 plants. Threats come from feral grazing animals, alien plant species, fire, and rats. This is the only Hawaiian species not formerly placed in Chamaesyce.

Euphorbia halemanui Sherff. ‘akoko, ‘ekoko, koko, spurge ENDEMIC | ENDANGERED

4.253 

Euphorbia herbstii

Small trees up to 8 m tall. The mostly narrowly oblong leaves are distichous and somewhat coriaceous. A rather large fruited species that is scattered in diverse mesic forests at elevations of 500–760 m in the Wai‘anae Mountains, O‘ahu. Populations have declined sharply in recent years, and in 2006 there were fewer than 60 mature individuals remaining. Threats have come from feral pigs and alien plant species, with fire as a potential threat. Listed in the Manual as Chamaesyce herbstii.

Euphorbia multiformis Hook. & Arn. ‘akoko, ‘ekoko, koko, spurge ENDEMIC | SECURE

4.250 

Euphorbia deppeana

Subshrubs with erect to decumbent stems to 120 cm in length. Leaves ovate to elliptic, distichous. Flowers solitary in the leaf axils. Rare, restricted to southern O‘ahu. Considered extinct until it was rediscovered in 1986 at the Nu‘uanu Pali, Ko‘olau Mountains. Known today from a single population with 50–100 plants. Threats come from alien plant species. Listed in the Manual as Chamaesyce deppeana.

4.252 

Euphorbia halemanui

4.254 

Euphorbia multiformis var. microphylla

99

100

FLOWERING PLANTS

Trees up to 9 m tall with elliptic to lanceolate leaves that are distichous. One of the tallest of the Hawaiian species of the Chamaesyce lineage. Once common, this plant is now much restricted in its distribution. Occurs in dry forest to subalpine forest at elevations of 600–2,800 m on West Maui and Hawai‘i. Listed in the Manual as Chamaesyce olowaluana.

are unique in the genus. Scattered in wet forest, usually along cloud-swept summit ridges at elevations of 600–830 m in the Ko‘olau Mountains, O‘ahu. The total number of extant plants is estimated to be between 200 and 400 individuals, with threats from alien plant species and feral pigs. Listed in the Manual as Chamaesyce rockii.

Euphorbia remyi A. Gray ex Boiss.

Euphorbia skottsbergii Sherff

ENDEMIC | RARE/EXTINCT

ENDEMIC | RARE/VULNERABLE/ENDANGERED

‘akoko, ‘ekoko, koko, spurge

4.255 

‘akoko, ‘ekoko, koko, spurge

Euphorbia multiformis var. multiformis

Shrubs with stems 0.5–2.0 m tall. Similar to and often confused with C. celastroides, from which it differs primarily in having a recurved fruit stalk. Leaves are distichous, typically 15–40 mm in length, and ovate to elliptic in shape. Scattered in open dry to wet forest at elevations of 200–500 m. The var. microphylla occurs on O‘ahu, Moloka‘i, West Maui, and leeward Hawai‘i, and var. multiformis occurs on O‘ahu and Maui. Listed in the Manual as Chamaesyce multiformis.

Euphorbia olowaluana Sherff ‘akoko, ‘ekoko, koko, spurge ENDEMIC | VULNERABLE

4.257 

Euphorbia remyi var. kauaiensis

Shrubs with erect stems to 2 m. The elliptic to oblong leaves are decussate. Three varieties of this species are recognized, all occurring on Kaua‘i, with the var. kauaiensis pictured. Occasional in wet forest and bogs at elevations of 150–900 m on Kaua‘i. This plant is quite variable throughout its distribution. Listed in the Manual as Chamaesyce remyi.

4.259 

Euphorbia skottsbergii var. skottsbergii

4.260 

Euphorbia skottsbergii var. vaccinioides

Euphorbia rockii C. N. Forbes ‘akoko, ‘ekoko, koko, spurge ENDEMIC | ENDANGERED

4.258 

4.256 

Euphorbia olowaluana

Euphorbia rockii

Compact shrubs or small trees to 2 m tall. The coriaceous, narrowly elliptic leaves are distichous. This plant is characterized by its large red capsules, which

Subshrubs with erect or ascending to prostrate stems to 1 m or more in length. The distichous leaves are variable in shape. An uncommon plant in coastal vegetation and dry shrubland but restricted to calcareous substrates at elevations of 5–200 m on O‘ahu, Moloka‘i, Maui, and Kaho‘olawe. The var. skottsbergii occurs on southwest O‘ahu and northeast Moloka‘i and is federally listed as endangered. The var. vaccini­ oides is on Moloka‘i, southern Maui, and Kaho‘olawe. Listed in the Manual as Chamaesyce skottsbergii.

FLOWERING PLANTS

Euphorbia sparsiflora A. Heller ‘akoko, ‘ekoko, koko, spurge ENDEMIC | RARE

4.261 

Euphorbia sparsiflora

Subshrubs with ascending stems to 60 cm tall. The elliptic to obovate leaves are decussate. Generally similar to Euphorbia multiformis, which has distichous leaves. Rare in the Wahiawa Bog on Kaua‘i at elevations of 600–800 m. Listed in the Manual as Chamaesyce sparsifolia.

Acacia, Vachellia, Prosopis, and Senna. In addition, there are many outstanding tropical ornamentals such as the jade vine (Strongylodon macrobotrys), shower trees (Cassia species and hybrids), flamboyant (Delonix regia), coral-tree (Erythrina species), and Colvillea and Bauhinia that are widely grown. Some species that are considered indigenous may have been brought in with early Polynesian colonization.

Acacia—A genus of about 1,075 species of shrubs and trees largely occurring in Australia. The name is from the Greek akis, sharp point, and this is in reference to the usually thorny nature of the plants. Some species have a symbiotic relationship with ants, such as the bullthorn acacia of Central America. Others are important sources of timber, fuel, forage, gums, and ornamentals. In the Hawaiian Islands, there are 2 endemic species, both having true leaves as juveniles and later the petioles becoming modified into phyllodes that function as leaves. Most of the African species formerly placed in Acacia have now been separated as Senegalia, while most of the neotropical species are now placed in the genus Vachellia.

Acacia koa A. Gray koa, koa‘oha

ENDEMIC | SECURE

Fabaceae

(PEA FAMILY)

As recognized here, a large family of trees, shrubs, vines, lianas, and herbs characterized by the fruit, a legume. Three subfamilies are recognized in which the flowers are either caesalpinaceous (slightly irregular), mimosaceous (regular, many stamens), or papilionaceous (irregular). There are 790 genera and about 19,500 species worldwide, making this the third largest family of flowering plants. Economically important for food, fodder, oils, resins, dyes, and ornamentals. In the Hawaiian Islands, there are 1 endemic, Kanaloa, and 11 indigenous genera: Acacia, Canavalia, Erythrina, Mezoneuron, Mucuna, Senna, Sesbania, Sophora, Strongylodon, Vicia, and Vigna. At least 39 other genera have naturalized, including tree species of

4.262 

Acacia koa

Trees up to 35 m tall, with variable morphology. Mature leaves are reduced to phyllodes that are narrowly elliptic and strongly sickle shaped. A dominant element of the vegetation in dry to wet forest at elevations of 60–2,060 m on all of the main islands except Ni‘ihau and Kaho‘olawe. This is the most valuable

timber tree in the Hawaiian Islands. The Hawaiians used the wood for making canoes, calabashes, and sometimes for the timbers of grass houses. It is highly prized for bowl making today.

Acacia koaia Hillebr. koai‘a

ENDEMIC | VULNERABLE

4.263 

Acacia koaia

Closely related to koa and sometimes considered to be conspecific. Koai‘a is usually distinguished by growing as a short tree up to 5 m tall with a broad and often gnarled crown. Phyllodes are relatively straight and narrow compared to the generally broader, more curved phyllodes of koa. Wood much denser and more fine grained than koa. Seed pods generally narrower than koa, with the seeds arranged longitudinally rather than transversely across the pod. Once common in dry forests on all of the main islands except Ni‘ihau, Kaho‘olawe, and perhaps O‘ahu. It is restricted in distri­ bution today and most common on the lower, south-facing slopes of Halea­ kalā in small populations or as isolated individuals.

Caesalpinia—A genus of trees, shrubs, and climbers. There are about 40 species, mostly from the Americas and Africa. The name honors Andrea Caesalpinio, a fourteenth-century Italian botanist and philosopher. Some species are important as timber sources, while others are cultivated as ornamentals, for dyes, tannins, or medical properties. Some species have become woody invaders outside of their natural range. In the Hawaiian Islands represented by 2 indigenous climbers. The Hawaiian species formerly known as Caesalpinia kavaiensis is now classified as Mezoneuron kavaiensis.

101

102

FLOWERING PLANTS

Caesalpinia bonduc (L.) Roxb.

Canavalia galeata (Gaudich.) Vogel

INDIGENOUS | SECURE

ENDEMIC | SECURE

hihikolo, kākalaioa, gray knickers

‘āwikiwiki, puakauhi

the species of the genus. In the Hawaiian Islands represented by the indigenous Entada phaseoloides, a huge woody liana. Known only from 2 collections on Kaua‘i (Powerline Trail) and last seen in 1920. Possibly a casual introduction.

Erythrina—A genus of about 120 species

4.265 

Canavalia galeata

Perennial climbers. Leaves pinnately trifoliolate, leaflets chartaceous, oblongovate to elliptic. Corolla dark purple. Pods 9.5–13.0 cm in length. Primarily in mesic forest, occasionally in alien vegetation such as guava and lantana shrubland on O‘ahu at elevations of 180–800 m.

Canavalia kauaiensis J. D. Sauer ‘āwikiwiki, puakauhi ENDEMIC | SECURE

4.264 

of trees and shrubs found throughout the tropics and warm temperate regions. The name is from the Greek erythros, red, because it is the predominant flower color in the genus. Many species are cultivated for their showy blooms, usually appearing before or with the formation of new leaves. In the Hawaiian Islands, there is 1 endemic species. Over 90 species and hybrids are cultivated in the Islands. The gall wasp, a nonnative introduction, has destroyed large numbers of both native and ornamental Erythrina.

Erythrina sandwicensis O. Deg. wiliwili

ENDEMIC | SECURE

Caesalpinia bonduc

Climbers to 30 m in length or scandent shrubs with recurved prickles on the young branches. The leaves are bipinnately compound and prickly on the rachis. Flowers yellow or upper one orange at base. Seeds can float for a long distance without losing their viability and thus are widely dispersed in pantropical oceanic islands. In the Hawaiian Islands in dry, mostly open, disturbed areas at elevations up to 230 m on Midway (Sand Island), Laysan, and all of the main islands except Lāna‘i and Kaho‘olawe. The seeds are used in making lei and are powdered for medicinal purposes. Children also used them as marbles.

Canavalia—A genus of about 60 species of pantropical lianas, vines, or perennial herbs. The name is from the common Indian name kanavali applied to the widespread C. maritima. There are 6 closely related endemic species that are perennial climbers and 2 naturalized species. The flowers of both the native Hawaiian species and the introduced species are used in the making of lei.

4.266 

Canavalia kauaiensis

Perennial climbers. Leaves pinnately trifoliolate, leaflets chartaceous, broadly elliptic to ovate. Corolla magenta to dark purple, with a white spot and streaks at the base. Pods 13–18 cm in length. Closely related to C. galeata. Occurs in diverse mesic forest and alien shrubland at elevations of 180–800 m on Kaua‘i.

4.267 

Erythrina sandwicensis

4.268 

Erythrina sandwicensis

Entada—A pantropical genus of 28 species

of lianas, shrubs, or trees. Leaves bipinnately compound. Most numerous in Africa and Madagascar. Name derived from the native Malabar name for one of

FLOWERING PLANTS

Deciduous trees up to 15 m tall with papery, reddish bark and a somewhat spiny trunk and branches. Flowering occurs during the dry season when the trees are leafless. Flower color varies from orange to yellow, white, or pale green. Locally common in dry forest up to 600 m elevation on all of the main islands. The soft wood is still used for outriggers of traditional Hawaiian canoes. Formerly the wood was also used for fishnet floats and surfboards. The red to orange seeds are strung in lei making. The erythrina gall wasp, which has caused defoliation and death of afflicted Erythrina trees, was first discovered in the Hawaiian Islands in Honolulu in April 2005 and rapidly dispersed throughout O‘ahu. Within six months, it had spread statewide and devastated nearly all of the susceptible native and cultivated species of Erythrina. The alarming rate of spread and impact of the erythrina gall wasp has put our native species at risk of future extinction. A biocontrol agent is now being used to combat the invader.

Kanaloa—A monotypic genus of woody legume shrub discovered in 1992 from 2 plants growing on Pu‘u Koa‘e, a steep, rocky sea stack just offshore of the ‘Ale‘ale area of Kaho‘olawe. Kanaloa shares a number of traits with members of the Dichrostachys and Leucaena groups but is distinct in its features from any other genus. Named for Kanaloa, a Hawaiian god, and an earlier name for Kaho‘olawe Island. According to Hawaiian legends, Kanaloa used the island as a place to rest and restore his energies. Pollen data from Kaua‘i, O‘ahu, and Maui suggest that the genus was once widespread in the Islands.

Kanaloa kahoolawensis Lorence & K. R. Wood kā palupalu o Kanaloa

ENDEMIC | ENDANGERED

4.270 

Kanaloa kahoolawensis

Woody shrub 0.5–1.0 m tall. Leaves divided into 3 broad leaflets, with a leaf nectary at the joint between each pair of leaflets. Inflorescences develop from the leaf axils with the flush of new leaves. The inflorescence is a globose head with 20 to 54 white flowers. Known today in the wild from only a single plant off southern Kaho‘olawe (Pu‘u Koa‘e) and in very limited cultivation.

Mezoneuron—A genus of 25–30 species of woody plants, formerly considered to be a subgenus of Caesalpinia. The center of distribution lies in Southeast Asia, with species in Africa, Australia, New Caledonia, Madagascar, and the Hawaiian Islands. The name comes from the Greek mezzo, greater, and neuron, nerve, referring to the winged pod. Mezoneuron is distinguished from Caesalpinia by its laterally compressed, indehiscent fruit with a winged upper suture. Represented in the Hawaiian Islands by a single endemic species.

Shrub or small tree 4–10 m tall, with stems lacking prickles. Corolla rose colored and showy. Pods with a longitudinal wing near their upper margin. An extremely rare species known only from populations in dry to mesic forest at 80–920 m elevation on Kaua‘i, the Wai‘anae Mountains on O‘ahu, and the North Kona District on Hawai‘i. Former populations on Lāna‘i and West Maui are now extinct. The Hawaiians are reported to have made spears from the very hard and durable wood. Fewer than 100 plants are known today. Reproduction is highly limited, possibly due to rat predation on the seeds.

Mucuna—A genus of 105 species of lianas, vines, erect herbs, or shrubs found throughout the tropics. The name is from the Brazilian common name, mucuna-quaca, for Mucuna urens, which is also present in the Hawaiian Islands. The legume pods of many species are covered with irritating hairs. In the Hawaiian Islands, there are 2 indigenous and 1 endemic species. The nonnative, red-flowered Mucuna bennetti is cultivated.

Mucuna gigantea (Willd.) DC. kā‘e‘e, kā‘e‘e‘e, sea bean INDIGENOUS | SECURE

Mezoneuron kavaiense (H. Mann) Hillebr. kāwa‘u, uhiuhi, kea

ENDEMIC | ENDANGERED

4.272 

4.269 

Kanaloa kahoolawensis—habit

4.271 

Mezoneuron kavaiense

Mucuna gigantea

Large woody lianas with stems to 15 m long with pinnately trifoliolate leaves. The pods are compressed and covered with irritating orange-brown hairs. Widespread from East Africa, India, China,

103

104

FLOWERING PLANTS

Malesia, and the Pacific Islands. Occurs sprawling over rocks, shrubs, and trees near the ocean or along streams at elevations up to 310 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i. The seeds were used medicinally by the Hawaiians as a violent cathartic and in the making of lei.

Sesbania tomentosa Hook. & Arn. ‘ohai

ENDEMIC | ENDANGERED

Senna—A genus of about 300 species of

herbs, shrubs, and trees mostly of the neotropics and subtropics and warm to cool temperate regions. Other species are found in Africa, Madagascar, Australia, Southeast Asia, and the Pacific Islands. These species were once included in Cassia. The name is from the Arabic, sanā. Many are grown as ornamentals and occasionally naturalizing in the Hawaiian Islands, such as the candle bush, Senna alata. There is 1 indigenous species and several cultivated species in the Hawaiian Islands.

4.274 

Sesbania tomentosa—Ka‘ena Point—habit

Senna gaudichaudii (Hook. & Arn.) H. S. Irwin & Barneby

Shrubs with decumbent or sprawling branches up to 4 m long or sometimes small trees to 6 m. The leaves have many silky smooth leaflets. Flower color varies from salmon to orange-red, scarlet, and yellow. Formerly occurring in lowland habitats but now mostly restricted to sandy beaches, dunes, and pond margins at elevations from sea level up to 830 m. On all of the main islands, as well as Necker and Nīhoa. A variable plant with respect to habit, flower color, and pubescence. Pictured here are plants from Ka‘ena Point (O‘ahu) and the distinctive variant form arborea from upland Moloka‘i. Although wide ranging, only about 2,000–3,000 plants remain, with the largest populations on Nīhoa. Loss of habitat and reduced population size have come with impacts of human development, offroad vehicle activity, feral grazing animals, alien plant species, and fire.

Sophora—A genus of about 50 species of shrubs and trees mostly of the tropics and subtropics. The name is of Arabic origin for an unknown member of this family. A few species are cultivated as ornamentals. In the Hawaiian Islands, there is 1 endemic species.

heuhiuhi, kolomona

INDIGENOUS | SECURE

4.275 

Sesbania tomentosa—Ka‘ena Point

Sophora chrysophylla (Salisb.) Seem. māmane, mamani

ENDEMIC | SECURE

4.273 

Senna gaudichaudii

Shrubs to 3 m tall. The flowers vary in color from greenish white to chartreuse or pale yellow. Widely dispersed in the Pacific Basin. In the Hawaiian Islands it is found in disturbed shrubland, dry forest, and the lower parts of mesic forest at elevations up to 920 m on all the main islands except Ni‘ihau.

4.276 

4.278 

Sesbania tomentosa—Moloka‘i

Morphologically polymorphic shrubs or small trees up to 15 m tall bearing leaves with many pinnate leaflets. The legume pods are moderately to strongly compressed between the seeds. In dry shrub­land and formerly in mesic to wet forest at elevations of 450–3,240 m on all of the main islands except Ni‘ihau and Kaho‘olawe. It is common in subalpine vegetation on East Maui and Hawai‘i. The wood was used by the Hawaiians for

Sesbania—A genus of trees, shrubs, and

perennial herbs represented by about 60 species that are found in the tropics and subtropics of the world. The name is from the Arabic sesban, which was used to designate Sesbania aegyptiaca. The Hawaiian Islands have 1 highly variable species and 1 introduced species. Some of the Hawaiian plants have horticultural merit.

4.277 

Sophora chrysophylla

Sesbania tomentosa—Moloka‘i—habit

FLOWERING PLANTS

sled runners and in the construction of houses, as handles for agricultural implements and adzes, and in certain religious ceremonies. Today it is used as fence posts because of its great durability.

Vicia menziesii Spreng.

Vigna marina (J. Burm.) Merr.

ENDEMIC | ENDANGERED

INDIGENOUS | SECURE

Hawaiian vetch

‘ōkolemakili, mohihihi, pūhili, beach pea

Strongylodon—A genus of 12 species of

vines and lianas found from Madagascar to Polynesia. The name is from the Greek strongylos, round, and odous, tooth, because of the rounded calyx teeth. The blue jade vine, Strongylodon macrobotrys, and the red-flowered S. nova-guianensis are cultivated in the Hawaiian Islands. There is 1 endemic species in the Hawaiian Islands.

4.281 

Vigna marina

Creeping or climbing perennial herbs up to several meters long. The leaves are pinnately trifoliolate. Pantropical in distribution. Occurs at the vegetation line on sandy beaches and occasionally inland at elevations up to 120 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i.

Strongylodon ruber Vogel kā ‘i‘iwi, nuku, nuku ‘i‘iwi ENDEMIC | RARE

Flacourtiaceae (FLACOURTIA

FAMILY)—Many genera of Flacourtiacae,

4.280 

4.279 

Strongylodon ruber

Lianas with herbaceous branches and pinnately trifoliolate leaves. The flowers may be in inflorescences up to 45 cm long. Found climbing in trees or shrubs in mesic to wet forest at elevations of 180– 825 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i.

Vicia—A genus of about 160 species of

annual or perennial herbs, often climbing with tendrils. Although the genus is primarily in temperate regions of North America, there are a few species distributed in South America and tropical East Africa. The name is from the Latin vicia used to designate these plants. Some species are cultivated for forage, for their edible seeds, or as ornamentals. In the Hawaiian Islands, there is 1 endemic species and at least 3 introduced species.

Vicia menziesii

Coarse, perennial climbing herbs that are supported in the canopy of trees and shrubs. The flowers, the largest in the genus, are yellowish white when first opening, turning to rose-purple with age. The species exhibits an insular phenomenon of gigantism with its large leaflets, flowers, and fruit. An extremely rare plant known from 2 populations on the island of Hawai‘i (Keauhou-Kīlauea, Pu‘uwa‘awa‘a) in wet to mesic forest at an elevation of about 1,600 m. This was the first Hawaiian plant to be federally listed as endangered. Today there are less than 50 plants in total.

Vigna—A pantropical genus of about 104

species of climbing or prostrate herbs or subshrubs, with a few species becoming erect herbs or small shrubs. The name honors Dominico Vigna, an Italian botanist of the seventeenth century. Some species are cultivated for forage, fiber, and edible seeds and pods. The Hawaiian Islands have 1 endemic and 2 indigenous species, one of which, Vigna adenantha, is considered to be extinct in the Hawaiian Islands, although it is extant elsewhere.

including Xylosma, which is present in the Hawaiian Islands, have now been placed in the Salicaceae.

Gentianaceae

(GENTIAN FAMILY)

Annual or perennial herbs, sometimes subshrubs, and rarely shrubs or even small trees. A few members are known to be saprophytes. Key features are opposite leaves, an absence of stipules, and a calyx and corolla fused into a short to well-developed tube. There are 85 genera and about 1,600 species, cosmopolitan in distribution, with a distribution in temperate, subtropical, and tropical mountains. Economically the family is of little importance, except that some genera, such as Gentiana and Exacum, are cultivated as ornamentals. In the Hawaiian Islands represented by the indigenous genus Schenkia and the introduced weedy Centaurium.

Schenkia—A genus of 5 species of annual

or biennial herbs found distributed in Australia, Eurasia, North Africa, and the Hawaiian Islands. The name honors Heinrich Schenck, a German botanist. In the Hawaiian Islands, there is 1 endemic species.

105

106

FLOWERING PLANTS

Schenkia sebaeoides Griseb. ‘āwiwi, centaury

ENDEMIC | ENDANGERED

Geranium—A genus of annual or perennial herbs or, in the Hawaiian Islands, shrubs and subshrubs. There are about 260 species found mostly in temperate regions but with some in tropical mountains. The name is from the Greek geranos, crane, because the beak of the fruit closely resembles the bill of a crane. In the Hawaiian Islands, there are 6 endemic species confined to the higher mountains of Kaua‘i, Maui, and Hawai‘i. These represent a single clade with close relatives in the western United States. They have sometimes been treated as a separate genus. Four other species have been introduced.

Geranium cuneatum Hook. hinahina, nohoanu ENDEMIC | SECURE

4.285 

Geranium cuneatum subsp. cuneatum

4.286 

Geraneum cuneatum subsp. hololeucum

4.287 

Geranium cuneatum subsp. tridens

Geranium arboreum A. Gray hinahina, nohoanu

ENDEMIC | ENDANGERED

4.282 

Schenkia sebaeoides

Erect annual herbs to 20 cm tall with somewhat fleshy leaves. The flowers are white or pale pink. A rare plant on volcanic or clay soil in dry rocky coastal sites on Kaua‘i, O‘ahu (extinct), Moloka‘i, Lāna‘i, and West Maui. As an annual plant, population size varies significantly between years. A census in 2010 found about 6,000 individuals, with the largest numbers on Moloka‘i. Threats come from alien woody species such as Casuarina equisetifolia and Prosopis pallida, as well as habitat degradation from livestock. Listed in the Manual as Centaurium sebaeoides.

4.283 

Geranium arboreum—habit

(GERANIUM FAMILY)

4.284 

Geranium arboreum

Annual or perennial herbs, subshrubs, or shrubs. Key features are the usually 5-merous flowers and the beaked or lobed fruit that splits into segments, with part of the style forming an awn or bristlelike appendage that is often coiled. There are 5 genera and about 650 species widespread in temperate and warm temperate regions and a few in montane tropics. Economically the family is of little importance. Geranium and Pelargonium are widely cultivated. In the Hawaiian Islands represented by the indigenous genus Geranium and the introduced Erodium.

Large shrubs to 4 m tall with 5–8-nerved ovate to cordate leaves. Uncommon in gulches in subalpine shrubland at elevations of 1,520–2,150 m on Haleakalā, East Maui. The irregular shape of the corolla is apparently an adaptation to bird pollination, which is otherwise unknown in the genus. This plant never becomes a true tree as the species epithet would indicate. Fewer than 500 plants are thought to remain in the native range. Threats come from cattle, feral pigs, and alien plant species.

Geraniaceae

Compact, erect, many-branched shrubs to 1 m tall. Often forms a dominant element of the plant cover in subalpine shrubland and forest and higher alpine shrubland, often as a pioneer on lava, at elevations of 1,480–3,250 m on East Maui and Hawai‘i. There are 4 named subspecies. The subsp. cuneatum is found at Humu‘ula, Mauna Kea, Hualālai, and South Kona on Hawai‘i, while subsp.

FLOWERING PLANTS

hololeucum has leaves that are densely grayish silky on both surfaces and occurs at Mauna Kea, Kahuku, ‘Āinahou, above Kūlani Prison, and Mauna Loa on Hawai‘i. The subsp. tridens, which occurs on Haleakalā, East Maui, has leaves that are densely silky-hairy on both surfaces and 3 apical teeth.

Geranium multiflorum A. Gray hinahina, nohoanu

ENDEMIC | ENDANGERED

Geranium hanaense A. C. Medeiros & H. St. John hinahina, nohoanu

Cyrtandra calpidicarpa (Rock) H. St. John ha‘iwale, kanawao ke‘oke‘o

ENDEMIC | VULNERABLE

ENDEMIC | SECURE

4.290 

4.288 

Geranium hanaense

Decumbent shrubs to 50 cm tall. Leaves silvery, with 3–5 apical teeth, similar to those of G. cuneatum. It forms large patches in bogs at an elevation of 1,670– 1,680 m in the Hāna Forest Reserve, East Maui. It is thought to have evolved from the high-elevation G. cuneatum in a manner similar to the 2 other bog Geranium species with dry site ancestors. Known from 2 bogs with 500–700 individuals.

Geranium hillebrandii Aedo & Muñoz Garm. ENDEMIC | RARE

4.289 

Geranium hillebrandii

Decumbent subshrub trailing in moss mats and other herbaceous vegetation of wet bogs. Branches rooting at the nodes, with leaves only at their ends. Rare in bogs at elevations of 1,490–1,770 m at Pu‘ukukui and Mount ‘Eke, West Maui. Known from 4 populations but abundant in number. Listed in the Manual as G. humile.

fertile staminal filaments. In the Hawaiian Islands, 59 endemic species are currently recognized, plus at least 67 hybrid combinations. These endemic species are thought to be derived from 3 colonizing lineages. This is one of the most polymophic and taxonomically complex genera in the Hawaiian Islands. The Manual described 53 species of Cyrtandra.

Geranium multiflorum

Compact, many-branched shrubs to 2 m tall. The leaves are green above and graygreen to gray-silky beneath. The flowers are white with purple streaks or pink to dark purple. A rare plant in montane grassland to wet forest or subalpine shrub­land at elevations of 1,580–2,450 m on Haleakalā, East Maui. Current estimates suggest that no more than 3,000 plants exist. Feral goats and pigs and alien plant species are the major threats.

Gesneriaceae

(AFRICAN VIOLET FAMILY)

Perennial herbs and shrubs (sometimes epiphytic), occasionally lianas, and rarely small trees. Key features are the often showy flowers, the leaves in pairs with one often reduced in size, and the fruit a capsule except in Cyrtandra, which has a berry. There are 139 genera and about 2,900 species that are pantropical except for a few found in temperate regions. Economically the family is of importance for the large number of cultivated genera, such as gloxinia (Sinningia), African violet (Saintpaulia), and Cape primrose (Streptocarpus). Other cultivated genera include Achimenes, Kohleria, Aeschynanthus, and Episcia, which are grown in the Hawaiian Islands.

Cyrtandra—A genus of about 600 species of perennial herbs, shrubs, or occasionally small trees found in Malesia and the Pacific. The highest concentration of species is in New Guinea and Borneo, but with a center of origin for Pacific species in the Fijian region. The name is from the Greek kyrtos, curved, and andros, stamens, because of the curved nature of the

4.291 

Cyrtandra calpidicarpa

Shrubs 1–4 m tall. The leaves in whorls of 3–4 in the upper nodes, elliptic to oblanceolate, 9–28 cm in length, upper surface largely glabrous, lower surface sparsely pubescent. Its unusual calyx is narrowly fusiform with a pronounced beak. The berries are curved, subcylindrical, and smooth. Occurs in disturbed mesic valleys at elevations of 120–600 m in the Ko‘olau Mountains, O‘ahu.

Cyrtandra cordifolia Gaudich.

ha‘iwale, kanawao ke‘oke‘o, hahala ENDEMIC | SECURE

4.292 

Cyrtandra cordifolia

Shrubs 1–7 m tall with branched stems. Leaves borne on the upper 3–8 nodes, heart shaped, 15–45 cm in length, moderately hairy on the upper surface and densely velvety-hairy beneath. Berries oval shaped, densely hairy. Occurs in moist,

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FLOWERING PLANTS

shaded gulch bottoms and slopes in disturbed mesic valleys at elevations of 270– 460 m in the Ko‘olau Mountains, O‘ahu.

Cyrtandra dentata H. St. John & Storey ha‘iwale, kanawao ke‘oke‘o ENDEMIC | ENDANGERED

Shrubs 1–4 m tall. Leaves borne on upper 3–6 nodes, with opposite paired leaves often unequal, chartaceous, broadly elliptic to ovate, 10–25 cm or more in length, upper surface sparsely pubescent, lower surface sparsely to densely velvety. Berries with scattered short hairs. Occurs in shaded mesic canyons and valleys and occasionally in wet forests at 300–850 m elevation in the Wai‘anae Mountains and leeward Ko‘olau Mountains (Wahiawā to Niu Valley), O‘ahu.

Shrubs 1.0–2.5 m tall with manybranched stems. Leaves borne on the upper 4–10 nodes, with opposite paired leaves often unequal, elliptic, 6–20 cm in length, chartaceous, leaf surfaces sparsely pubescent, raised veins on lower surface. Occurs in wet or rarely mesic forest at elevations of 530–1,900 m on Maui.

Cyrtandra lessoniana Gaudich. ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

Cyrtandra grandiflora Gaudich. ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

4.293 

Cyrtandra dentata

Shrubs 1.5–5.0 m tall, stems sparingly branched. Leaves borne on upper 3–4 nodes, with opposite paired leaves often unequal, broadly elliptic to ovate, 9–33 cm in length, chartaceous, upper surface sparsely pubescent, lower surface sparsely to densely velvety. Berries ovoid, densely hairy. Occurs in shaded canyons in moist valleys at elevations of 300–600 m in the northern Wai‘anae Mountains and leeward northern Ko‘olau Mountains (Pa‘ala‘a, Kawailoa divide to ‘Ōpae‘ula), O‘ahu. Populations have increased in recent years and there are now 6 known populations with more than 1,600 plants. Threats come from feral pigs and alien plant species.

Cyrtandra garnotiana Gaudich.

hahala, ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

4.295 

Cyrtandra grandiflora

Shrubs 1–2 m tall with few-branched stems. Leaves borne on the upper 3–8 nodes, with opposite paired leaves often unequal, elliptic to ovate, 7–36 cm in length, mostly smooth above and hairy along the veins below, veins slightly raised on both surfaces, chartaceous. The berries are smooth and ellipsoid. Occurs in wet canyon bottoms or lower slopes of disturbed mesic valleys and wet forest at elevations of 300–540 m in the Ko‘olau Mountains, O‘ahu.

Cyrtandra hashimotoi Rock ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

4.294 

Cyrtandra garnotiana

4.296 

Cyrtandra hashimotoi

4.297 

Cyrtandra lessoniana

Shrubs 2–4 m tall with many-branched stems. Leaves borne on the upper 3–7 nodes, with opposite paired leaves often unequal, lanceolate to oblanceolate, 5–21 cm in length, upper surface with scattered hairs, lower surface with dense velvety pubescence, veins strongly impressed above. The berries are oval shaped and smooth. Occurs in mesic to wet forest at elevations of 365–1,230 m in the Wai‘anae and Ko‘olau Mountains on O‘ahu.

Cyrtandra lysiosepala (A. Gray) C. B. Clark ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

4.298 

Cyrtandra lysiosepala

FLOWERING PLANTS

Shrubs 3–5 m tall with few-branched stems. Leaves borne on the upper 3–6 nodes, with opposite paired leaves often unequal, chartaceous, elliptic, 13–23 cm in length, lightly hairy on the upper surface and hairy on the veins beneath. Calyx lobes in this species are almost leaflike. Berries ellipsoid, smooth. Occurs in wet forest at elevations of 950–1,430 m on leeward side of Hawai‘i.

Cyrtandra macrocalyx Hillebr. ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

Low, decumbent, succulent subshrubs with stems with adventitious roots and becoming subterranean. Leaves fleshy, elliptic to ovate, 10–31 cm in length. Flowers totally hidden by the leaves. Berries cylindrical. Rare in wet forest along mossy sites or rocks near waterfalls at elevations of 540–800 m on Kaua‘i (KīlaueaHa‘ena, Wahiawa-Hanapēpē, Olokele Canyon). Population size for the 3 areas of occurrence totals less than 500 individuals. Threats come from feral pigs and alien plant species.

Cyrtandra platyphylla A. Gray

‘ilihia, ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

Cyrtandra paludosa Gaudich.

hahala, moa, ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

4.302 

4.299 

Cyrtandra platyphylla

Shrubs up to 3.5 m tall with unbranched to few-branched stems. Leaves borne opposite to occurring in whorls of 3–4 on the upper 6–22 nodes, well spaced, suborbicular to broadly ovate, 4–34 cm in length, moderately to densely hairy above, densely velvety hairy along the veins. Berries ellipsoid, smooth, and surrounded by the persistent calyx. Occurs in wet to sometimes mesic forest in shaded sites at elevations of 370–1,525 m on Maui and Hawai‘i.

Cyrtandra macrocalyx

Shrubs 1–2 m tall with few-branched stems. Leaves borne on the upper 3–5 nodes, in whorls of 3–4, narrowly elliptic, 7–16 cm in length, hairy on the upper surface and velvety short-hairy beneath. Calyx large and leaflike. Berries oval shaped and smooth. A rare species from streamsides and gulches in mesic valleys and wet forest at elevations of 915–1,130 on Moloka‘i (Waikolu to Wailau valleys, Kaunakakai Gulch) and West Maui (‘Īao Valley).

Cyrtandra procera Hillebr.

ha‘iwale, kanawao ke‘oke‘o ENDEMIC | SECURE

Cyrtandra oenobarba H. Mann ha‘iwale, kanawao ke‘oke‘o ENDEMIC | RARE

4.301 

4.300 

Cyrtandra oenobarba

Cyrtandra paludosa var. paludosa

Shrubs 1–5 m tall with unbranched or few-branched stems. Leaves somewhat fleshy, elliptic lanceolate, smooth, with impressed veins above and nearly smooth beneath. Berries ellipsoid, smooth. The var. paludosa is common in diverse mesic forest to wet forest at elevations of 240–1,280 m on Kaua‘i, O‘ahu, Maui, and Hawai‘i, while the var. microcarpa is restricted to Kaua‘i.

4.303 

Cyrtandra procera

Shrubs 1–2 m tall with mostly unbranched stems. Leaves in whorls of 4–6 on the upper 5–10 nodes, elliptic, 5.5–17.0 cm in length, smooth and shiny above and short velvety-hairy beneath. The

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FLOWERING PLANTS

oval-shaped berries are glabrous. Occurs in wet forest at elevations of 1,130–1,270 m on Moloka‘i (Waikolu to Pelekunu valleys, Pēpē‘ōpae Bog).

Cyrtandra propinqua C. N. Forbes

valleys at elevations of 350–520 m on the leeward side of the Ko‘olau mountains (Pauoa to Mānoa valleys), O‘ahu.

Cyrtandra spathulata H. St. John ENDEMIC | SECURE

ENDEMIC | SECURE

Shrubs 1–7 m tall with branched stems. Leaves borne on the upper 3–6 nodes, with opposite paired leaves often unequal, broadly elliptic to suborbicular, 9–27 cm in length, sparsely hairy above and moderately velvety hairy beneath. Berries oval shaped, densely hairy. Occurs in moist shaded gulch bottoms in disturbed mesic valleys to diverse mesic and wet forest at elevations of 460–1,100 m in the Wai‘anae Mountains, O‘ahu (Mākua Valley to Palikea).

Cyrtandra wawrae C. B. Clarke ENDEMIC | SECURE

4.304 

Cyrtandra propinqua

Shrubs 2.5–7.0 m tall, stems branched. Leaves opposite, borne on the upper 3–5 nodes, chartaceous, very broadly elliptic to very broadly ovate, 15–30 cm in length, upper surface somewhat rugose, lower surface with raised veins. Berries ellipsoid, glabrous. Occurs in moist, shaded gulch bottoms and slopes in disturbed mesic valleys and wet forest at elevations of 210–610 m in the northern Ko‘olau Mountains (Kaunala to Kīpapa and Waiāhole gulches), O‘ahu.

Cyrtandra sandwicensis (H. Lév.) H. St. John & Storey ENDEMIC | VULNERABLE

4.306 

Cyrtandra spathulata

Shrubs 2–3 m tall, unbranched to few branched. Leaves borne on the upper 3–8 nodes, with opposite paired leaves often unequal, elliptic to ovate, 10–25 cm in length, chartaceous, upper surface with sparse hairs, lower surface with moderate density of velvety hairs, veins slightly raised on both surfaces. Berries ovoid. Occurs in wet forests at elevations of 275– 1,590 m on Maui.

Cyrtandra waianaeensis H. St. John & Storey ha‘iwale, kanawao ke‘oke‘o, hahala ENDEMIC | SECURE

4.308 

Cyrtandra wawrae

Small shrubs 0.5–2.0 m tall, stems with few branches. Leaves borne on the upper 2–5 nodes, with opposite paired leaves often unequal, peltate to broadly ovate or suborbicular, 24–41 cm in length, sparse to moderate pubescence on the upper surface, moderately velvety hairy below. Berries subglobose. Occurs infrequently in mesic forest at elevations of 120–620 on Kaua‘i.

Goodeniaceae

(GOODENIA FAMILY)

4.305 

Perennial herbs, shrubs, or small trees. Key features are the strongly irregular flowers and the presence of a pollen-collecting cup subtending the stigmas. There are 11 genera and about 400 species found primarily in Australasia. Economically the family is of no importance except for the cultivation of a few species of Scaevola. In the Hawaiian Islands represented by the indigenous genus Scaevola.

Cyrtandra sandwicensis

Shrubs 0.5–1.0 m tall, stems unbranched. Leaves borne on the upper 3–8 nodes, with opposite paired leaves often unequal, elliptic to ovate, 6.5–20.0 cm in length, chartaceous, moderately to sparsely pubescent above, densely velvety hairy below. Berries ovoid with scattered short hairs. Occurs on moist slopes and in shaded bottoms of gulches in disturbed

4.307 

Cyrtandra waianaeensis

Scaevola—A genus of perennial herbs, shrubs, or small trees with 97 species primarily from Australia, with 2 widely distributed species on tropical beaches. The name is from the Greek scaevous, lefthanded or awkward, probably referring

FLOWERING PLANTS

to the shape of the flower, which looks as though it has been torn in half. In the Hawaiian Islands, there are 9 endemic species and 1 indigenous species. Scaevola hobdyi from West Maui is now considered to be extinct. The Hawaiian species represent 3 distinct colonizing lineages. Scaevola glabra and S. taccada form 2 of these lineages, while the other 7 species together form the third lineage. These 3 lineages represent half of the 6 lineages of Scaevola that extend outside of Australia The “half-flowers” of Scaevola have been immortalized in old Hawaiian legends.

Scaevola chamissoniana Gaudich.

Prostrate perennial herbs, woody only at the base of older stems. Leaves succulent, obovate to spatulate, 2–5 cm in length. Flowers 1–3 in axillary cymes, corolla yellowish green externally, cream colored inside. Formerly occurred in coastal sites on all of the main islands except Kaho‘olawe. Currently only 1 significant population exists on stabilized coastal dunes at Waiehu, West Maui, plus relictual populations from Kaupō, Maui, and on small islets off Maui and Moloka‘i. Total population has been estimated at less than 300 individuals. Threats come from feral grazing animals and alien plant species.

naupaka kuahiwi

Scaevola gaudichaudiana Cham.

ENDEMIC | SECURE

ENDEMIC | SECURE

Shrubs to 1 m tall. Leaves oblanceolate, 3–7 cm in length. Flowers 1–3 in axillary cymes, corolla yellowish brown or pinkish, fragrant. Occurs on dry ridges and flats in open shrubland and forest at elevations of 75–800 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Scaevola glabra Hook. & Arn.

‘ohe naupaka

ENDEMIC | SECURE

4.313 

4.311 

4.309 

Scaevola chamissoniana

Shrubs or sometimes small trees to 2.5 m tall; leaves oblanceolate, 9–14 cm in length. Flowers 7–15 in open cymes, corolla white, often with purple lines on the interior, very fragrant. Occurs in wet forest and open areas at an elevation of about 350 m on Moloka‘i, Lāna‘i, Maui, and Hawai‘i.

Scaevola gaudichaudiana

Shrubs or occasionally small trees, 1–3 m tall. Leaves broadly oblanceolate to ovate, 5–10 cm in length, coriaceous. Flowers 7–18 or more in open cymes, corolla white, glabrous. Occurs in wet forest and open clearings at 170–800 m elevation on Kaua‘i and O‘ahu.

Scaevola gaudichaudii Hook. & Arn. naupaka kuahiwi

ENDEMIC | SECURE

Scaevola glabra

Shrubs 1.5–4.0 m tall, occasionally epiphytic. Leaves coriaceous, clustered at the ends of the branches, oblanceolate to obovate, 5–15 cm in length. Flowers 1 or rarely 2 in the leaf axils, corolla bright yellow. Occurs in wet forest at elevations of 200–800 m on Kaua‘i and in the Ko‘olau Mountains, O‘ahu. The yellow, bird-pollinated flowers are unlike any other Scaevola in the Hawaiian Islands. This character and its tetraploid nature indicate that this species represents a lineage of colonization distinct from other Hawaiian species. The flowers in the Kaua‘i populations are larger than those found on O‘ahu.

Scaevola kilaueae O. Deg.

huahekili uka, naupaka kuahiwi, pāpa‘ahekili ENDEMIC | RARE

Scaevola coriacea Nutt. dwarf naupaka

ENDEMIC | ENDANGERED

4.314 

4.310 

Scaevola coriacea

4.312 

Scaevola gaudichaudii

Scaevola kilaueae

Shrubs to 1 m tall. Leaves coriaceous, oblanceolate, 3–7 cm in length, clustered at the tips of the stems. Flowers 3–7 in

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FLOWERING PLANTS

cymes, corolla purplish brown, lobes cream with purple lines, and throat pale yellow-brown. Occurs locally on old lava flows and ash substrate in open ‘ōhi‘a forest and shrubland at elevations of 1,000– 1,460 m in the Ka‘ū District and vicinity of Kīlauea in Hawai‘i Volcanoes National Park.

Scaevola mollis Hook. & Arn. naupaka kuahiwi

Small trees or shrubs, 1.5–3.5 m tall. Leaves oblanceolate, 4–10 cm in length. Flowers 3 in cymes, crowded in the upper branches, corolla white with purple lines inside. Occurs in wet forest at elevations of 700–1,400 m on Kaua‘i and Moloka‘i.

Scaevola taccada (Gaertn.) Roxb.

naupaka kahakai, huahekili, naupaka, naupaka kai INDIGENOUS | SECURE

ENDEMIC | SECURE

on steep rock faces. A few species are occasionally cultivated.

Gunnera—A genus of 40 species of predominantly Southern Hemisphere herbs, often growing in wet places. These are moderate to giant herbs with large orbicular to peltate leaves. The name honors Johan Ernst Gunnerus, an eighteenth-century Norwegian botanist. The peeled petioles are eaten in Chile. In the Hawaiian Islands, there are 2 endemic species usually found growing on wet cliff faces in cloud-swept forests.

Gunnera petaloidea Gaudich. ‘ape‘ape

ENDEMIC | SECURE

4.315 

Scaevola mollis

Large shrubs to 2.5 m tall. Leaves lanceolate to oblanceolate, 5–15 cm in length, lower surface covered with soft gray hairs. Flowers 3 in cymes, corolla purple or rarely white. Occurs in wet forest on ridges, valleys, and in boggy areas at elevations of 600–1,400 m on Kaua‘i and O‘ahu, where it is common, and on Moloka‘i, where it is rare. Hybrids are common between this species and S. gaudichaudiana.

Scaevola procera Hillebr. ENDEMIC | SECURE

4.317 

Scaevola taccada

Shrubs up to 3 m tall. Leaves are smooth or slightly hairy, 5–20 cm in length, somewhat variable in shape from oblanceolate to suborbicular, usually with inrolled margins. Flowers in cymes in the upper leaf axils, corolla pale yellowish green, fruit white, compressed globose in shape. A common plant throughout tropical and subtropical Pacific and Indian ocean coasts. In the Hawaiian Islands in coastal sites throughout the archipelago, except absent from Gardner Pinnacles, Necker, and Nīhoa. Commonly cultivated, especially around hotels and condominiums along the coast. Listed in the Manual as Scaevola sericea.

Gunneraceae

(GUNNERA FAMILY)

4.316 

Scaevola procera

Large terrestrial perennial herbs with often gigantic leaves. Comprises only a single genus with about 40 species extending from Mexico to Chile, tropical and South Africa, Malesia, Tasmania, subantarctic islands, and the Hawaiian Islands. The rhizomes exhibit a symbiotic relationship with the cyanobacterial genus Nostoc. Nitrogen fixation with this relationship aids the plants and allows them in grow in nitrogen-poor soils and

4.318 

Gunnera petaloidea

Giant herbs from 1 to 6 m tall with large leaves up to 90 cm wide and 100 cm in length. It can be distinguished from G. kauaiensis by leaves not peltate. The perfect or unisexual flowers are tiny and numerous on the elongated inflorescence. Fruit yellow to orange. Occurs on steep slopes in cloud-swept wet forest at elevations of 920–1,300 m or higher on Kaua‘i (Blue Hole), O‘ahu, eastern Moloka‘i, Maui, and the Kohala Mountains of Hawai‘i.

Hydrangeaceae

(HYDRANGEA FAMILY)

Shrubs, small trees, lianas, and sometimes subshrubs and rhizomatous perennial herbs. Sometimes included with the Saxifragraceae but distinct from it, especially in the woody nature of most of the genera. There are 17 genera and about 220 species that are found in temperate and subtropical regions of the Northern Hemisphere, with a few representatives

FLOWERING PLANTS

in Southeast Asia, Malesia, and South America. Economically the family is important horticulturally for such genera as Deutzia, Hydrangea, and Philadelphus. In the Hawaiian Islands represented by the genus Broussaisia.

Broussaisia—An endemic Hawaiian genus

of 1 species; unisexual flowers borne on separate plants. Distinctive from other genera in the family by its berrylike fruit and the styles joined together. The name honors François-Joseph-Victor Broussais, an eighteenth-century French philosopher and physiologist. The genus is a sister group to a clade from southern China and Malesia. Recent studies suggest that Broussaisia is best considered within the genus Hydrangea, and a future placement in that genus is likely.

Broussaisia arguta Gaudich. kanawao

ENDEMIC | SECURE

4.319 

Broussaisia arguta—female

Shrubs up to 5 m tall. Leaves opposite or in whorls of 3, coriaceous, and with the upper surface having deeply impressed veins. Plants dioecious. The female flowers have no petals, while those of the males have petals that may be green, pink, or purplish blue. The dark red fruit is readily eaten by birds. A common wet forest plant at elevations of 400–1,400 m or more on all of the main islands except Ni‘ihau and Kaho‘olawe.

Broussaisia arguta—male

Haplostachys haplostachya (A. Gray) H. St. John honohono

ENDEMIC | ENDANGERED

Hydrophyllaceae (WATERLEAF FAM-

ILY)—The former Hydrophyllaceae, represented in the Hawaiian Islands by the indigenous genus Nama, has now been merged into the Boraginaceae.

Lamiaceae

(MINT FAMILY)

Annual or perennial herbs, occasionally vines, shrubs, or even small trees. Key features are the presence of aromatic oils (absent in Hawaiian species), usually quadrangular stems with opposite leaves, a verticillate inflorescence, and a 4-lobed ovary. As newly redefined to include much of the Verbenaceae, the family includes 238 genera and about 6,500 species. These are cosmopolitan in distribution, with the Mediterranean and Central Asia being centers of diversity. Economically the family is important as a source of essential oils, culinary herbs, ornamentals, and medicinals. They include catnip (Nepeta), thyme (Thymus), basil (Ocimum), oregano (Origanum), rosemary (Rosmarinus), lavender (Lavandula), horehound (Marrubium), mint (Mentha), sage (Salvia), coleus (Plectanthus), and others. In the Hawaiian Islands represented by 2 endemic genera, Stenogyne and Haplostachys, and 3 indigenous genera: Phyllostegia, Plectranthus, and Lepechinia. Stenogyne, Haplostachys, and Phyllostegia represent a single founding lineage related to Stachys from North America. At least 10 other genera of mints have become naturalized in the Islands.

Haplostachys—An endemic Hawaiian genus

4.320 

at each axil of the inflorescence. Flowers 2 per whorl, nutlets dry. The plants are usually nonaromatic.

of 5 species of erect perennial herbs somewhat woody at the base. Only 1 species is extant, with the others extinct. The name is from the Greek haplo, single, and stachys, spike, since only 1 flower appears

4.321 

Haplostachys haplostachya

Erect perennial herbs with dense white hairs on stems. The narrowly heartshaped leaves are green above with dense white hairs beneath. Formerly known from Kaua‘i and Maui and now extant at Kīpuka Kālawamauna, Pu‘ukapele, and Waili‘i on Hawai‘i. Although highly restricted in distribution today, close to 20,000 plants remain. Threats include alien plant species, human disturbance, and fire.

Lepechinia—A genus of about 40 species of aromatic subshrubs from western South America, Mexico, California, and the Hawaiian Islands. The name honors Ivan Lepechin, an eighteenth-century Russian botanist. Some species have horticultural merit. In the Hawaiian Islands represented by a single indigenous species.

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FLOWERING PLANTS

Lepechinia hastata (A. Gray) Epling pakaha, pitcher sage

Phyllostegia ambigua (A. Gray). Hillebr. ENDEMIC | SECURE

INDIGENOUS | SECURE

Lepechinia hastata

Erect perennial herbs to 1.5 m tall. The sagittate leaves are hairy on both surfaces and aromatic. Native to Baja California and some of the islands off Baja California, Socorro Island, and presumably native to Maui. Occurs on Haleakalā in open shrubland and pastures at elevations of 610–1,220 m. This species is possibly an early introduction to the Hawaiian Islands rather than a product of long-distance dispersal, as whaling ships visited both Baja California and the Hawaiian Islands.

Phyllostegia—A genus of 34 species of nonaromatic herbs, lianas, or shrubs with 32 species endemic to the Hawaiian Islands; the others occur in Tahiti and Tonga. The name is from the Greek phyllon, leaf, and stego, cover, probably referring to the calyx lobes, which are leaflike and enclose the flower in the type species. Foliage not aromatic. Flowers terminal, with the upper corolla lip typically conspicuously shorter than the lower lip. Fruit a fleshy drupe. Many species are sympatric, but hybridization does not appear to be common. The Manual included only 27 species.

kāpana

ENDEMIC | SECURE

4.323 

4.322 

Phyllostegia grandiflora (Gaudich.) Benth.

Phyllostegia ambigua

Scandent subshrubs or lianas, stems densely to sparsely hairy. Leaves somewhat thick and chartaceous or relatively thin and membranaceous, ovate to elliptic, 11–16 cm in length. Flowers 6–16 per whorl, corolla white, occasionally tinged with pink. A highly variable species in leaf morphology, pubescence, and flower structure. Occurs in wet forest at elevations of 915–1,980 m on Moloka‘i (Kapulei), Maui, and Hawai‘i (Mauna Loa).

4.325 

Phyllostegia grandiflora

Lianas or scandent subshrubs, stems densely covered with long hairs. Leaves somewhat coriaceous, 9–16 cm in length, pubescent on both surfaces. Flowers fragrant, 6–18 per whorl, corolla white. Common in mesic to wet forest, especially cloud-swept summits, at elevations of 300–1,230 m in the Ko‘olau and Wai‘anae mountains (Mount Ka‘ala), O‘ahu.

Phyllostegia hirsuta Benth. ENDEMIC | ENDANGERED

Phyllostegia floribunda Benth. ENDEMIC | RARE

4.324 

Phyllostegia floribunda

Erect subshrubs with stems 40–90 cm tall. Leaves membranaceous and rather limp, somewhat hairy, variable in shape from ovate to lanceolate, 12–24 cm in length. Flowers 2 per whorl, corolla dark red, white at base. Occurs in mesic to wet forest at elevations of 430–1,130 m on Hawai‘i (Honokai‘a, Pāhala, Kohala Mountains, Kīlauea to Laupāhoehoe). Known from 12 populations with fewer than 100 individuals.

4.326 

Phyllostegia hirsuta

Erect subshrubs to scandent lianas, stems densely hairy. Leaves rugose, ovate, 7.3– 18.0 cm in length, with dense pubescence on both surfaces. Flowers 6–16 per whorl, corolla white, tinged with purple on the

FLOWERING PLANTS

upper lip. Occurs in diverse mesic forest to wet forest at elevations of 300–1,070 m in the Ko‘olau and central Wai‘anae mountains, O‘ahu. Known from 9 populations with about 160 mature plants. Threats come from feral pigs and goats and alien plant species.

Phyllostegia lantanoides Sherff ENDEMIC | SECURE

Scandent vines, stems densely hairy. Leaves ovate, 15–24 cm in length, pubescent on both surfaces. Flowers 10–20 per whorl in congested inflorescences, corolla white, tinged with pink. Occurs along streambeds and shaded places in wet forest at elevations of 365–1,350 m on Hawai‘i (Hīlea to Laupāhoehoe and Kohala Mountains).

Plectranthus—A genus of about 200 species of herbs and subshrubs, often succulent, found in paleotropical and subtropical regions, especially Africa. The name is from the Greek plectron, spur, and anthos, flower, referring to the spurlike protuberance on the upper side of the corolla. Some species are cultivated for their edible tubers, medicinal properties, or as ornamentals. In the Hawaiian Islands, there is 1 indigenous species.

Plectranthus parviflorus Willd. ‘ala‘ala wai nui, spurflower INDIGENOUS | SECURE

4.327 

Phyllostegia lantanoides

Scandent to decumbent lianas, stems with short pubescence. Leaves membranous, ovate to lanceolate leaves, 4–9 cm in length, rugose on the upper surface, both sides with small glandular dots. Flowers typically 4 per whorl, corolla white, tinged with purple. Occurs in wet forest at elevations of 640–1,190 m in the Ko‘olau and central Wai‘anae mountains, O‘ahu.

Phyllostegia vestita Benth. ENDEMIC | SECURE

4.328 

Phyllostegia vestita

4.329 

Plectranthus parviflorus

Stenogyne angustifolia A. Gray ENDEMIC | ENDANGERED

4.330 

Stenogyne angustifolia

Decumbent to scandent vines, stems weakly 4 angled. Leaves narrowly ovate to lanceolate, 2.6–5.3 cm in length. Flowers 2 per whorl, yellow to dull brownish pink, magenta, or red. Formerly known from dry subalpine shrubland at elevations of 1,550–2,150 m on Moloka‘i and Maui but presently occurring today only in the Pōhakuloa Training Area on Hawai‘i. Although thousands of plants remain, these are localized in a fairly small area. Threats come from fire, feral pigs, other ungulates, and alien plant species.

Stenogyne calaminthoides A. Gray ENDEMIC | SECURE

Erect or ascending perennial herbs from a fleshy tuberous base. The ovate or roundish leaves are hairy on both surfaces, with small oil globules on the upper surface. Native from Australia to Malesia and Polynesia. In the Hawaiian Islands occurs on dry, exposed sites on all of the main islands except Kaho‘olawe.

Stenogyne—An endemic Hawaiian genus of nonaromatic perennial vines or erect herbs. There are 22 recognized species, and Maui has the greatest diversity, with 10 species. Distinguished by axillary flowers and the upper lip of the corolla usually conspiculously longer than the lower lip. The fleshy fruit of this genus and Phyllostegia is unusual in the family. The name is from the Greek stenos, narrow, and gyneo, gyneocium.

4.331 

Stenogyne calaminthoides

115

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FLOWERING PLANTS

Scandent or decumbent vines up to several m long, stems sharply to weakly 4 angled. Leaves membranous, ovate, 2.2–5.1 cm in length. Typically 6 flowers per whorl, corolla pink, dark magenta, purple, or rarely white. A relatively common species in mesic to wet forest and low, boggy forest at elevations of 750–1,550 m on Hawai‘i.

Scandent or decumbent vines up to 1–2 m long, stems 4 angled. Leaves membranous, broadly ovate, 3.2–5.9 cm in length. Flowers 4–6 per whorl, corolla cream, pink, or red. Occurs in wet forests at elevations of 1,050–1,450 m on Moloka‘i and Maui. On Maui the flowers are pink or cream in color, and on Moloka‘i they are red.

Stenogyne kaalae Wawra

Stenogyne kanehoana O. Deg. & Sherff

ENDEMIC | SECURE/ENDANGERED

ENDEMIC | ENDANGERED

Trailing or scandent vines, stems weakly 4 angled. This species differs from S. purpurea, within which it was once included, primarily by its coriaceous leaves that are weakly revolute near the margins, with the margins subentire to weakly serrate or crenulate-serrate in the upper half, sometimes nearly to the base, and the lower surface glabrous. Rare in wet forest at 1,090–1,250 m elevation in Kōke‘e State Park, Kaua‘i. Known today from 5 populations with no more than 200 individuals. Threats come from feral pigs and other ungulates and alien plant species. Federally listed as endangered. Listed in the Manual as Stenogyne purpurea var. kealiae.

Stenogyne microphylla Benth. ENDEMIC | SECURE

4.334 

4.332 

Stenogyne kaalae subsp. kaalae

Scandent vines climbing up to 2 m, stem 4 angled. Leaves somewhat coriaceous, ovate-lanceolate, 4.2–7.7 cm in length. Flowers in whorls of 2–6, corolla dark maroon. Occurs in diverse mesic forest at elevations of 750–1,100 m on O‘ahu. Two subspecies are recognized, with subsp. kaalae from the Wai‘anae Mountains and a single collection from the Ko‘olau Mountains. The rare subsp. sherffii is known only from the Pe‘ahināi‘a Trail in the Ko‘olau Mountains.

Stenogyne kanehoana

Scandent vines with climbing stems 1–2 m in length. Leaves membranaceous, somewhat flaccid, densely hairy, 6–14 cm in length. Flowers 3–6 per whorl, corolla white or pale yellow with a pinkish purple lip. Extremely rare in diverse mesic forest at 700–750 m in the Wai‘anae Mountains, O‘ahu (Pu‘ukānehoa). Known from only 2 individuals when declared endangered in 2003, and these have since died. One additional individual discovered in the following year may still exist. Threats have been from feral pigs, alien plant species, and military training. The species survives in cultivation.

Stenogyne kealiae Wawra ENDEMIC | RARE

Stenogyne kamehamehae Wawra ENDEMIC | SECURE

4.336 

4.333 

Stenogyne kamehamehae

4.335 

Stenogyne kealiae

Stenogyne microphylla

Scandent vines with stems up to several m in length, becoming woody at the base, 4 angled. Leaves small, ovate to elliptic, 0.4–1.3 cm in length. Flowers in whorls of 2, corolla greenish yellow to brownish pink. Occurs in subalpine forest at elevations of 1,200–2,700 m on East Maui and Hawai‘i.

FLOWERING PLANTS

Stenogyne purpurea H. Mann ENDEMIC | SECURE

Trailing vines or erect herbs, 20–70 cm tall, spreading by rhizomes, stems terete or 4 angled. Leaves membranous, variable in shape from ovate to lanceolate, variably glabrous or densely hairy on both surfaces. Flowers 2–16 per whorl, corolla typically dark to dull red. Occurs in mesic forest and bordering lava flows at lower elevations and in subalpine woodland at elevations of 500–3,100 m on East Maui and Hawai‘i. An extremely variable plant that has the greatest elevational range of any species in the genus.

Vitex rotundifolia L. f.

kolokolo kahakai, pohinahina, beach vitex INDIGENOUS | SECURE

Stenogyne sessilis Benth. ENDEMIC | SECURE

4.340  4.337 

Low, branching shrubs with procumbent stems that often root at the nodes. The rounded leaves are aromatic when crushed and are densely white hairy beneath. Widespread from China and Japan south to Malesia, India, Mauritius, Australia, and the Pacific Islands. In the Hawaiian Islands a common strand plant on sandy beaches, shores, and dunes on all of the main islands except Kaho‘olawe. It was used medicinally in the past and is currently a popular cultivated plant in beach areas.

Stenogyne purpurea

Trailing or scandent vines, occasionally forming large mats. Stems weakly 4 angled, young stems with soft, downy hairs. Leaves membranous, the margin flat and conspicuously serrate or crenulate-serrate, 3.5–13.0 cm in length, lower surface usually at least sparsely appressed with soft, downy hairs. Flowers 2–6 per whorl, corolla pinkish purple or rarely white. Occurs in diverse mesic to wet forest at elevations of 600–1,300 m on Kaua‘i.

Stenogyne rugosa Benth. mā‘ohi‘ohi

ENDEMIC | SECURE

4.338 

Stenogyne rugosa

Vitex rotundifolia

4.339 

Stenogyne sessilis

Scandent or trailing vines, sometimes forming a dense ground cover, stem sharply 4 angled. Leaves membranous, broadly ovate, 1.9–6.5 cm in length, petioles virtually absent. Flowers typically 2 per whorl, corolla white, pink, or dark purple and cream inside. Occurs in mesic to occasionally wet forest at elevations of 900–1,700 m on Lāna‘i (now extinct), Maui, and Hawai‘i.

Vitex—A genus of trees and shrubs, sometimes vines, with about 250 species found primarily in the tropics and subtropics but with some representation in temperate Europe and Asia. This genus was formerly placed in the Verbenaceae. The name is from the Latin viere, to twist, as some of the species have flexible stems. In the Hawaiian Islands represented by 1 widespread indigenous species. Other species are cultivated in the Islands.

Lauraceae

(LAUREL FAMILY)

Trees and shrubs and the parasitic vine Cassytha. Key features are small flowers with an undifferentiated perianth, the stamens usually in 4 whorls of 3, and the fruit a drupe with a single seed. There are 52 genera and about 2,550 species found in the tropics and warm regions, especially in Southeast Asia, Malesia, and Brazil. Economically the family is important as a source of timber, aromatic oils, spices, and food, especially the avocado. In the Hawaiian Islands represented by 2 indigenous genera, Cassytha and Cryptocarya.

Cassytha—A genus of 23 species of the Old

World and the New World, especially Australia. One pantropical species occurs in the Hawaiian Islands. The name is from the Aramaic kesatha, a tangled wisp of hair, referring to the filiform vining stems. All the species are parasitic.

117

118

FLOWERING PLANTS

Cassytha filiformis L.

kauna‘oa mālolo, kauna‘oa pehu INDIGENOUS | SECURE

4.341 

Cassytha filiformis

Pantropical parasitic vines on a number of different host plants. Stems very thin and green to orange in color. Leaves reduced to minute scales. In the Hawaiian Islands, mostly in coastal sites on Kure Atoll and all of the main islands except Kaho‘olawe.

Cryptocarya—A genus of about 300 species of trees from the tropics, especially tropical Asia, but absent from central Africa. The name is from the Greek krypto, hidden, and karyon, nut, referring to the floral tube that encloses the fruit. In the Hawaiian Islands represented by 1 endemic species.

Loganiaceae

(LOGANIA FAMILY)

Trees, shrubs, lianas, and occasionally herbs. Key features are opposite leaves with stipules usually present, symmetrical flowers and a 2-chambered superior ovary. Flowers may be perfect or functionally unisexual. There are 14 genera and about 400 species that are predominantly in tropical and subtropical regions and rarely temperate areas. The family is of little economic importance except for a few cultivated genera such as Gelsemium and Spigelia. Curare and strychnine are obtained from species of Strychnos. In the Hawaiian Islands represented by the endemic genus Labordia.

Labordia—An endemic Hawaiian genus

of 17 species of shrubs and small trees, all thought to be derived from a single colonizing lineage. Plants functionally dioecious. The name honors M. Laborde, a marine officer on the voyage of the French exploratory ship Uranie. The genus is closely related to Geniostoma found across the Pacific Basin.

Labordia hedyosmifolia Baill. kāmakahala

ENDEMIC | SECURE

Labordia hirtella H. Mann. kāmakahala

ENDEMIC | SECURE

4.344 

Labordia hirtella

Shrubs or small trees to 6 m tall with winged or sharply angled stems. Leaves glossy, elliptic to obovate, 6–19 cm in length. Flowers in sessile cymes that are open to congested in arrangement, corolla color varies from orange to yellow or even cream, capsules ellipsoid to ovoid and slightly compressed, 15–34 mm in length. Scattered in mesic to wet forest at elevations of 500–1,830 m on eastern Kaua‘i, O‘ahu (Ko‘olau Mountains), Moloka‘i, Lāna‘i, Maui, and Hawai‘i.

Labordia hosakana (Sherff) Wagner, Herbst & Sohmer kāmakahala ENDEMIC | RARE

Cryptocarya mannii Hillebr. hōlio

ENDEMIC | SECURE

4.343 

4.342 

Cryptocarya mannii

Trees 6–10 m tall and up to 30–35 cm in diameter. Branches angular and with a gray, corky outer layer when mature. The alternately arrayed coriaceous leaves are 6–15 cm in length, with a yellow midrib. An uncommon tree in open stands of dry to mesic forest at elevations of about 900– 1,200 m on Kaua‘i and O‘ahu.

Labordia hedyosmifolia

Terrestial (rarely epiphytic), sprawling, or erect branched shrubs to shrubby trees to 4.5 m tall, sometimes with scandent and liana-like stems angled to weakly winged. Highly variable plants with respect to leaf shape and habit. Leaves narrowly to broadly elliptic to ovate, 5.0–13.5 cm in length. Flowers in sessile or subsessile cymes, corolla orangish yellow, capsules ellipsoid to ovoid, 12–20 mm in length. Scattered to locally common in wet forest but also bog margins and sometimes mesic forest at elevations of 700–1,590 m on Moloka‘i, Lāna‘i, Maui, and Hawai‘i.

4.345 

Labordia hosakana

Low-growing shrubs up to 0.5 m tall with angled stems. Leaves coriaceous with impressed veins on the upper surface, variable in shape, 1.5–5.0 cm in length. Flowers in sessile cymes, corolla yellowish orange, capsules broadly ovoid, 10–13 mm in length. Occurs on cloud-swept summits in wet forest at elevations of 790–860 m in the Ko‘olau Mountains, O‘ahu (Kīpapa to Wa‘ahila).

FLOWERING PLANTS

Labordia kaalae C. N. Forbes kāmakahala

ENDEMIC | RARE

fruits. The var. tinifolia, the most widespread of these, is pictured. The vulnerable var. lanaiensis occurs on Lāna‘i, while the endangered var. wahiawaensis is restricted to Wahiawa Valley on Kaua‘i.

Labordia venosa Sherff kāmakahala

ENDEMIC | SECURE

4.346 

Labordia kaalae

Shrub or small tree 2–4 m tall or occasionally up to 6 m, stems terete. Leaves membranous, broadly to lanceolate elliptic, 6–17 cm in length, lower leaf surface and young stems with moderate to light covering of short hairs. Flowers in open cymes with peduncles that elongate with fruit maturity, corolla green to yellowish green, capsules ovoid, 12–20 mm in length. Rare in mesic and sometimes wet forest at elevations of 450–950 m in the Wai‘anae Mountains, O‘ahu.

Labordia tinifolia A. Gray kāmakahala

ENDEMIC | SECURE/VULNERABLE/ENDANGERED

Labordia waiolani Wawra kāmakahala

ENDEMIC | SECURE

4.348 

Labordia venosa

Shrubs to 2.5 m tall, stems angled and many branched. Leaves thin, coriaceous, oblanceolate to elliptic, 4–15 cm in length, veins strongly impressed on the upper surface and conspicuously raised on the under surface. Flowers in sessile cymes, corolla yellow to yellow orange, capsules ovoid, 12–20 mm in length. Occurs in wet forest at elevations of 910–1,910 m on East Maui.

Labordia waialealae Wawra

kāmakahala, kāmakahala lau li‘i ENDEMIC | SECURE

4.347 

4.350 

Labordia waiolani

Branched shrubs 1–3 m tall, stems angled to slightly winged. Leaves glossy, narrowly elliptic to elliptic, 4–12 cm in length, upper surface with veins slightly impressed. Flowers in sessile cymes, corolla dark to golden yellow, capsules ellipsoid, 11–17 mm in length. Occurs in mesic to wet forest at elevations of 360– 1,280 m on O‘ahu, Moloka‘i, and rarely on Lāna‘i.

Malvaceae

Labordia tinifolia var. tinifolia

Shrubs or small trees 2–8 m tall, stems dichotomously branched, terete. Leaves membranous, elliptic to lanceolate, 4.5–21.0 cm in length. Flowers in open pendulous inflorescences, corolla pale yellow-green, capsules ovoid, 8–17 mm in length. Occurs on ridges, slopes, or understory in mesic to wet forest at elevations of 300–920 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Three varieties are recognized that basically differ in the size of the flowers and

Compact or sometimes openly branched shrubs to 2 m tall, stems angled, many branched. There are 2 conspicuous forms of this plant. One is a compact shrub with small leaves less than 4 cm in length clustered at the branch tips. The other is an openly branched shrub with larger leaves distributed along the branches. Flowers in subsessile cymes, corolla yellow, capsules ovoid, 9–15 mm in length. Occurs in wet forest and shrubby bog hummocks at elevations of 1,220–1,560 m on Kaua‘i (Mount Wai‘ale‘ale, Alaka‘i Swamp, Pihea, Makaleha Mountains) and possibly Moloka‘i (Kamalō).

(MALLOW FAMILY)

4.349 

Labordia waialealae

Herbs, shrubs, and trees. Key features are the usual presence of stellate hairs, often glandular, the often palmately lobed or divided leaves, and large spiny pollen grains. The family has recently been expanded based on molecular data to now include the Bombacaceae, Sterculiaceae, and Tiliaceae. Under this concept, there are 113 genera and about 5,000 species that are cosmopolitan in distribution but with centers of diversity in the neotropics. Economically the family is

119

120

FLOWERING PLANTS

important for a number of ornamentals such as Hibiscus, hollyhocks (Althaea), turk’s cap (Malvaviscus), and mallows (Malva). Okra is the fruit of Hibiscus esculentus. Cotton comes from the seed capsules of Gossypium. In the Hawaiian Islands represented by the endemic genera Hibiscadelphus and Kokia and the indigenous Abutilon, Gossypium, Hibiscus, and Sida. Thespesia is questionably indigenous. Weedy species of Malva, Modiola, Malachra, Malvastrum, and Sidastrum have become naturalized.

Very rare in dry forest on Lāna‘i (Kahulu and Maunalei valleys, Kānepu‘u Ridge, Kehewai Gulch) at elevations of 210–520 m. Known only from a single population of no more than 100 individuals. Threats come from grazing by mouflon sheep and deer and alien plant species.

Abutilon incanum (Link) Sweet

‘ilima pua kea, ko‘oloa kea, ma‘o, sweet hoary abutilon INDIGENOUS | SECURE

Abutilon—A genus of about 160 species of herbs, shrubs, and small trees found primarily in the tropics and subtropics, especially the neotropics but also Australia. The name is from the Greek word, probably of Arabic origin, designating plants in this family. In the Hawaiian Islands, there are 3 endemic species, 1 indigenous, and 3 naturalized. Each native species represents a separate colonizing lineage. A few species are grown as ornamentals and are known as flowering maples.

Abutilon eremitopetalum Caum ENDEMIC | ENDANGERED

Abutilon sandwicense (O. Deg.) Christoph. ENDEMIC | ENDANGERED

4.352 

Abutilon incanum

Subshrubs or shrubs up to 1 m tall, densely white or silvery stellate hairs on the stems. Leaf blades cordate-ovate, 1–6 cm in length, margins serrate to crenate, petioles half or more of the length of the blade. Flowers white to pink or yellow with maroon veins. Only the pink-flowered forms are found in the Hawaiian Islands. Native to the Southwestern United States and Mexico, but also in the Hawaiian Islands in dry habitats at elevations from sea level up to 220 m on all of the main islands except Hawai‘i. This range suggests that it may have been a historical introduction.

Abutilon menziesii Seem. ko‘oloa ‘ula

ENDEMIC | ENDANGERED

4.351 

Abutilon eremitopetalum

Shrubs with grayish stems with stellate glandular. Leaf blades heart shaped, 7–12 cm in length, margins crenate-dentate, petioles as long as the blades. Flowers solitary or paired in the leaf axils. The tiny petals are included within the calyx.

Diffusely branched shrubs up to 2 m tall with dense velvety stellate hairs on the stems. Leaves heart shaped, 3–10 cm in length. Flowers deep maroon (East Maui and Hawai‘i) or pale red (Lāna‘i). Rare and local in dry forest at elevations of 200–520 m on O‘ahu(?), Maui, Lāna‘i, and Hawai‘i. Although widespread in distribution, no more than 10 populations with 450–500 individuals remain. Threats come from agricultural and urban development, livestock grazing, goats, and competition with alien plant species. An attractive species suitable for cultivation.

4.353 

Abutilon menziesii

4.354 

Abutilon sandwicense

Shrubs up to 3 m or more tall, with dense stellate hairs on the stems. Leaves heart shaped, 8–22 cm in length. Flowers bright green to reddish brown with green veins, solitary and pendulous. Rare on steep slopes in dry forest at elevations of 300–600 m in the Wai‘anae Mountains, O‘ahu (between Makaleha Valley and Pu‘ukaua). Known currently from less than 20 populations totaling 300–500 individuals. Threats come from feral pigs and goats, cattle grazing, and alien plant species.

Gossypium—A genus of 49 species of herbs, subshrubs, shrubs, or small trees found in tropical and subtropical regions around the world. The name is from the

FLOWERING PLANTS

Greek gossypion, cotton. The Hawaiian Islands have 1 endemic and 2 naturalized species. Cotton was grown commercially in the Islands for nearly a century but never achieved any importance as an industry.

Hibiscadelphus distans L. E. Bishop & D. R. Herbst hau kuahiwi ENDEMIC | ENDANGERED

Gossypium tomemtosum Nutt. ex Seem. huluhulu, ma‘o, Hawaiian cotton ENDEMIC | VULNERABLE

Trees up to 7 m tall and with diameters up to 30 cm. Leaves broadly oval in shape usually with 3 (5) main veins, 10–25 cm in length, both surfaces pubescent in the vein angles. Flowers solitary, dark magenta. Once present at Kīpukapuaulu on the eastern slopes of Mauna Kea at an elevation of 1,310 m. Last observed in the wild in 1930 and now extinct, although surviving in cultivation. Hybrids between this species and H. hualalaiensis were once common at Kīpukakī, Hawai‘i, where both species were planted, but they have since been destroyed.

Hibiscadelphus hualalaiensis Rock hau kuahiwi

ENDEMIC | EXTINCT

4.356 

4.355 

Gossypium tomentosum

Shrubs 0.5–1.5 m tall, softly white tomentose. Leaf blades wider than long, 3–10 cm in width, 3–5 lobed. Flowers solitary or in a few short cymes, petals bright yellow. The seeds in the capsule are embedded in reddish brown lint. Occurs primarily in arid, rocky, or clay coastal plains at elevations from sea level to 120 m on all of the main islands except Hawai‘i. This species has not achieved economic importance. The introduced sea island cotton (G. barbadense) has naturalized in the Islands. A notable feature of G. tomentosum is the absence of extrafloral nectaries, the only species in the genus without this trait. This has been hypothesized to result from the absence of native ants in the Hawaiian Islands, and thus there was no selective pressure for a coevolved defense system.

Hibiscadelphus—An endemic genus of 7 species of shrubs or trees, with 6 species thought to be extinct in the wild and the other one extremely rare. The name means the brother of Hibiscus, because the 2 genera are closely related. The curved corollas are an adaptation to bird pollination, rare in the family. Three species survive in limited cultivation.

Hibiscadelphus distans

Shrubs or small trees up to 5 m tall and 5–8 cm in diameter, with a rounded crown and leaf blades broadly ovate, 4–10 cm in length, upper surface sparsely pubescent, more densely so on lower surface. Flowers solitary, erect, bright green at first fading to maroon. Very rare. Known originally from a single population discovered in 1972 in a remnant of dry forest above Koai‘e Stream in Waimea Canyon, Kaua‘i, at an elevation of about 350 m; it was later destroyed by a landslide. Known today from 2 populations with an estimated 20 wild and 150 reintroduced trees. Another population of 50 trees in the Hipalau Valley was destroyed in 1992 by Hurricane ‘Iniki.

Hibiscadelphus giffardianus Rock hau kuahiwi

ENDEMIC | EXTINCT

4.357 

Hibiscadelphus giffardianus

4.358 

Hibiscadelphus hualalaiensis

Trees to 7 m tall and 30 cm in diameter. Leaf blades ovate to rounded, 10–15 cm in length, usually with 3–5 main veins. Flowers 1–2 in the leaf axils, yellowish green fading to purple. A rare plant that once occurred in dry to mesic forest at Hualālai and Waihou, North Kona, Hawai‘i. The last known tree died in 1992 and the species is now considered to be extinct in the wild, although surviving in cultivation.

Hibiscus—A large genus of about 650 species of shrubs, trees, and occasionally herbs found in the tropics and subtropics of the world. The name is from the Greek hibiscos, mallow. In the Hawaiian Islands, there are 5 endemic, 2 indigenous, and 2 naturalized species. The native species are thought to represent at least 4 separate colonizing lineages. The hibiscus is the state flower of Hawai‘i. Many species, including the native ones, are in cultivation.

121

122

FLOWERING PLANTS

Hibiscus arnottianus A. Gray

aloalo, hau hele, koki‘o ke‘oke‘o, pāmakani ENDEMIC | SECURE/ENDANGERED

a maroon spot at their base, drying greenish purple. Rare and local in dry forest and shrubland at elevations of 130–800 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Three subspecies are recognized, with subsp. molokaiana from Moloka‘i. The pictured subsp. brackenridgei occurs on Lāna‘i, Maui, and Hawai‘i, while the subsp. mokuleianus is on Kaua‘i and O‘ahu. A plant cultivated because of its showy yellow flowers and selected as Hawai‘i’s state flower.

Hibiscus clayi O. Deg. & I. Deg. 4.359 

Hibiscus arnottianus subsp. arnottianus

Shrubs or small trees to 8 m tall, glabrous or young stems with yellow stellate pubescence. Leaf blades coriaceous, elliptic-ovate to broadly ovate, 4–15 cm or more in length, smooth to pubescent beneath. Flowers solitary near the ends of branches, white, weakly fragrant. Three subspecies are recognized. The subsp. arnottianus has a red staminal column and occurs in mesic to wet forest at elevations of 120–790 m in both the Wai‘anae and eastern Ko‘olau mountains on O‘ahu. The endangered subsp. immaculatus has a white staminal column and is very rare on Moloka‘i (Wailau, Waihānau, and Pāpalaua valleys). The subsp. punaluuensis with pubescent leaves, stems, and calyces occurs in wet forest at elevations of 200–700 m in the Ko‘olau Mountains (Kaipapa‘u to Waiāhole).

Hibiscus brackenridgei A. Gray aloalo, ma‘o hau hele

ENDEMIC | ENDANGERED/EXTINCT

koki‘o ‘ula

ENDEMIC | ENDANGERED

Perennial herbs or subshrubs 1.0–2.5 m tall. Stems covered with stellate hairs. Leaf blades ovate to orbicular, 5–15 cm in length, both surfaces with stellate pubescence. Flowers solitary in the upper leaf axils, pale magenta to rose, never open very widely. Distributed in the West Indies, Florida, Central America, and South America. In the Hawaiian Islands in wet disturbed areas at elevations of 90–240 m on Kaua‘i, O‘ahu, Maui, and Hawai‘i. Apparently it was once much more common than today.

Hibiscus kokio Hillebr. ex Wawra

aloalo, koki‘o, kokio ‘ula, kokio ‘ula ‘ula ENDEMIC | RARE

4.361 

Hibiscus clayi

Shrubs or trees 4–8 m tall, stems with stellate pubescence. Leaf blades shiny elliptic, 3–7 cm in length, glabrous. Flowers solitary, borne near the ends of branches, dark red. Closely related to the more common Hibiscus kokio. Occurs in dry forest at elevations of 230–350 m in eastern Kaua‘i (Nounou Mountain, Hāli‘i Valley, Anahola Mountains). A rare species known today from a single wild population with 4 individuals. Widely grown in cultivation.

4.363 

Hibiscus kokio subsp. kokio—habit

4.364 

Hibiscus kokio subsp. kokio

Hibiscus furcellatus Desr.

‘akiahala, ‘akiohala, hau hele, aloalo, hau hele wai INDIGENOUS | SECURE

4.360 

Hibiscus brackenridgei subsp. molokaiana

Sprawling to erect shrubs 1–3 m tall or sometimes small trees 5 m or more tall. Leaf blades roundish, 5–15 cm in length, 3, 5, or 7 lobed. Flowers solitary or few in short terminal racemes, petals yellow with

4.362 

Hibiscus furcellatus

FLOWERING PLANTS

rare tree known from only a few individuals, the last tree in the wild died in 1918. Marginally persisting in cultivation, where it has been grafted onto Kokia kauaiensis in botanical gardens.

Kokia drynarioides (Seem.) Lewton koki‘o, hau hele ‘ula

ENDEMIC | ENDANGERED

4.365 

Hibiscus kokio subsp. saintjohnianus

Shrubs or small trees 3–7 m tall. Leaf blades coriaceous, glossy green, elliptic to ovate, 3–10 cm in length, glabrous. Flowers solitary, borne near the end of branches, red, orange, or rarely yellow. The subsp. kokio with red flowers occurs in dry to wet forest at elevations of 70– 890 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and presumably Hawai‘i. The subsp. saintjohnianus with orange flowers is known only from dry to mesic forest in northwestern Kaua‘i.

Hibiscus waimeae A. Heller

aloalo, koki‘o ke‘oke‘o, koki‘o kea

ENDEMIC | SECURE/ENDANGERED

4.367 

Hibiscus waimeae subsp. waimeae

Trees, 6–10 m tall, with gray bark. Leaf blades rounded to ovate-elliptic. Leaves 5–18 cm in length, lower surface densely grayish velvety hairy. Flowers solitary, borne near the end of branches, strongly fragrant, white when first open fading to pink later in the day. Two subspecies are recognized. The subsp. waimeae with larger flowers that occurs in species-rich mesic forest at elevations of 250–1,200 m on Kaua‘i (Waimea Canyon, west and southwest valleys) is pictured. The subsp. hannerae from northwest Kaua‘i (Hanakāpī‘ai, Limahuli, Kalihi Wai) is endangered.

Kokia—An endemic Hawaiian genus of 4 species of trees with 1 extinct and the others rare. The name is from the Hawaiian name for these plants, koki‘o. One of the extant species persists only in cultivation. All 3 are cultivated for their showy red flowers resembling hibiscus.

Kokia cookei O. Deg.

4.369 

Kokia drynarioides

Trees up to 8 m tall. Leaf blades 12–25 cm in width, 7–9 lobed, lower surface nearly pubescent near its base and at the apex of the petioles. Flowers scarlet red. Rare in dry forest at elevations of 460–900 m at Pu‘uwa‘awa‘a and Hu‘ehu‘e in the North Kona District, Hawai‘i. Almost extinct in the wild but in limited cultivation. A 1998 survey found only 10 living trees fragmented in 3 small populations. Threats have come from cattle grazing, seed predation by rodents, and alien plant species.

Kokia kauaiensis (Rock) O. Deg. & Duvel koki‘o, hau hele ‘ula

ENDEMIC | ENDANGERED

koki‘o, hau hele ‘ula ENDEMIC | EXTINCT

4.370  4.368 

4.366 

Hibiscus waimeae subsp. waimeae—habit

Kokia cookei

Small trees 3–5 m tall, extinct in the wild. Leaf blades 5–13 cm in width, 5–7 lobed. Flowers orangish red. Formerly found in the Mauna Loa area on the arid western end of Moloka‘i. Always an extremely

Kokia kauaiensis

Trees 5–10 m tall. Leaf blades 12–25 cm in width, usually 7–9 lobed, lower surface nearly pubescent near its base and at the apex of the petioles. Flowers brick red. Of the extant species, this has the largest flowers. Occurs in diverse mesic forest at elevations of 350–660 m in western

123

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FLOWERING PLANTS

Kaua‘i (Pa‘aiki, Ku‘ia, Mahanaloa, Kalalau, and Koai‘e valleys). The population of this rare species once shrank to a single tree but with assistance has now increased to 3 small populations totaling about 145 individuals. Threats have come from goats and deer and alien plant species.

is from the Greek thespesios, divine, because the pantropical Thespesia populnea was collected in Tahiti on Cook’s voyage of 1769 where it had been planted about places of worship. In the Hawaiian Islands, the same species is now considered indigenous, although it may have been introduced by the Polynesians.

Sida—A genus of about 250 species of perennial herbs and subshrubs found primarily in tropical and subtropical areas. The name was used by Linnaeus from the writings of Theophrastus but for an unrelated plant. The Hawaiian Islands have 1 indigenous and 6 naturalized species, most of them weedy. Petals from the native species are used in making lei.

Thespesia populnea (L.) Sol. ex Corrêa

Waltheria indica L.

‘ala‘ala pū loa, ‘uhaloa, hala ‘uhaloa, hi‘aloa, kanakaloa INDIGENOUS | SECURE

milo, portia tree

INDIGENOUS/POLYNESIAN INTRODUCTION  |  SECURE

Sida fallax Walp. ‘ilima

INDIGENOUS | SECURE

4.373 

4.371 

Sida fallax

Prostrate or diffuse to erect spreading shrubs, stems 0.2–1.5 m in length, typically with dense stellate pubescence. Leaf blades lanceolate-ovate to suborbicular, 1–12 cm in length, bright green above and densely hairy beneath. Flowers axillary or solitary with 2–7 per node, corolla yellow to orange-yellow. Widespread on Pacific Islands westward to China. In the Hawaiian Islands, in coastal habitats, dry to diverse mesic forest, and occasionally low-elevation wet forest, with an overall elevational range from sea level to 1,950 m. On Midway Atoll, Nīhoa, and all of the main islands. A highly variable plant that probably represents a species complex not fully understood. The flowers are used in the making of lei and were formerly reserved for chiefs.

Thespesia—A genus of 17 species of shrubs or trees found in the South Pacific, Asia, Africa, and the Caribbean. The name

4.372 

Thespesia populnea

Arborescent tree 5–10 m or more in height. Leaf blades deltate to orbiculate, 5–30 cm in length, shiny yellow-green. Large yellow-colored flowers with a purple spot at the base of the petals, quickly deciduous. Pantropical in coastal habitats but thought to be native only to the paleotropics. Widely occurring in the Hawaiian Islands, often in cultivation along beaches, on all of the main islands except Kaho‘olawe. It remains unclear whether this species is indigenous or an early Polynesian introduction to the Hawaiian Islands.

Waltheria—A genus of 50–60 species of mostly subshrubs that have a range centered in the neotropics. This genus would formerly have been placed in the Sterculiaceae. The name honors A. F. Walther, an eighteenth-century professor at Leipzig. In the Hawaiian Islands represented by 1 indigenous pantropical species, which is often weedy.

Waltheria indica

Subshrub to 2 m tall covered with velvety yellow stellate hairs. The small yellow flowers are fragrant. Pantropical in distribution. In the Hawaiian Islands apparently indigenous in dry, often disturbed places at elevations from sea level up to 1,220 m on Midway Atoll and all of the main islands. The plant had medicinal value to the Hawaiians, who used the bitter inner bark and roots as a painkiller for sore throat. Formerly placed in the Sterculiaceae, a family now merged into the Malvaceae.

Menispermaceae (MOONSEED FAMILY)

Primarily herbaceous vines or lianas, rarely shrubs or small trees. Key features are the presence of male and female flowers on separate plants, the viny habit, a double whorl of sepals, and the usually 3-merous floral parts. There are 71 genera and about 450 species found primarily in the tropics and subtropics of the world, with a few species extending into temperate areas. Economically the family is of little importance except for a few genera that are cultivated, such as Cocculus and

FLOWERING PLANTS

Menispermum. In the Hawaiian Islands represented by the indigenous genus Cocculus.

Cocculus—A genus of 8 species of vines,

shrubs, or small trees found in Asia, Malesia, Africa, the Pacific, and North and Central America, but absent from South America and Australia. The name is from the Greek kokkos, berry, referring to the fruit. In the Hawaiian Islands, there is 1 indigenous species that is variable throughout its range.

Cocculus orbiculatus (L.) DC. huehue, hue

INDIGENOUS | SECURE

A number of species of Ficus are cultivated as ornamentals, and Ficus microcarpa has escaped to become a serious invasive species in disturbed native forests. In the Hawaiian Islands represented by the indigenous genus Streblus. Species of Ficus, Broussonetia, and Morus have become naturalized.

Streblus—A genus of about 14 species

of trees found in Africa, Malesia, eastern Australia, New Zealand, the Society Islands, Fiji, and the Hawaiian Islands. Flowers perfect or unisexual and then either monoecious or dioecious. The name is from the Latin streblus, flesh about the haunches, as the base of the fruit is often thick and fleshy. In the Hawaiian Islands, there is 1 indigenous species.

Streblus pendulinus (Endl.) F. Muell. a‘ia‘i

INDIGENOUS | SECURE

4.374 

Cocculus orbiculatus

Scandent or sprawling vines with variable-shaped leaves usually 3 nerved from the base. Fruits are dark blue at maturity. Extends from Southeast Asia to the Himalayas, Malesia, and across the Pacific. In the Hawaiian Islands occurs in open areas, on lava and lava crevices, and in dry to mesic forest at elevations from sea level to 1,400 m on all of the main islands except Kaho‘olawe. A variable species throughout its range. Listed in the Manual as Cocculus trilobus.

Moraceae

(MULBERRY FAMILY)

Trees, shrubs, and lianas. Key features are the presence of milky sap, male and female flowers on the same or different plants, and the usual presence of 2 stigmas. There are 38 genera and about 1,150 species found primarily in tropics and subtropics of the world. Economically the family is important for edible fruits such as figs (Ficus), mulberries (Morus), and breadfruit (Artocarpus).

4.375 

Streblus pendulinus

Shrubs or small trees 2–8 m tall, bark pale gray, stems with milky sap, and plants typically dioecious. Leaf dark green on upper surface, glossy, thin, chartaceous, shape variable from lanceolate to ovate, 6–13 cm in length. Flowers unisexual and plants typically dioecious, staminate spikes 4–11 cm in length. Occurs from New Guinea to Micronesia, south to Norfolk Island, eastern Australia, Fiji, and Rapa Nui (Easter Island). In the Hawaiian Islands mostly in mesic forest at elevations of 300–1,680 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Myrsinaceae

(MYRSINE FAMILY)—

The Myrsinaceae, including the Hawaiian genera Embelia and Myrsine, have now been merged into the Primulaceae.

Myrtaceae

(MYRTLE FAMILY)

Trees and shrubs. Typical features are the opposite, glandular punctate leaves, 5-merous petals (although the petals are minute or absent in several genera), and numerous and often brightly colored and conspicuous stamens. There are about 131 genera and some 5,500 species, mostly of tropical and subtropical areas but prevalent in temperate and subtropical Australia. Economically the family is important for its edible fruits (guavas, rose apples, and Surinam cherries), spices (allspice, cloves, and bay rum), timber (Eucalyptus), and cultivated plants (e.g., eucalyptus, bottlebrush, paperbark, and myrtles). In the Hawaiian Islands represented by the indigenous genera Eugenia, Metrosideros, and Syzygium.

Eugenia—A genus of about 950 species of trees and shrubs found mostly in the neotropics but with good representation in Africa and Malesia. The name honors seventeenth-century Prince Eugene of France. Some species are cultivated, such as the Surinam cherry (E. uniflora). In the Hawaiian Islands represented by 1 endangered endemic species and 1 indigenous species.

Eugenia reinwardtiana (Blume) DC. nīoi

INDIGENOUS | SECURE

Myoporaceae (MYOPORUM FAMILY)—

The Myoporaceae, including the Hawaiian genus Myoporum, have now been merged into the Scrophulariaceae.

4.376 

Eugenia reinwardtiana

125

126

FLOWERING PLANTS

Small trees or shrubs 1.5–7.0 m tall, young branches with light covering of brown pubescence. Leaves coriaceous, ovate to elliptic, 2.5–8.0 cm in length, upper surface glossy. Flowers 1–2 in leaf axils, petals 4. Fruit oval berries, red, orange, or yellow at maturity. Widely distributed in the Pacific from Malesia to Micronesia and the Tuamotus. In the Hawaiian Islands, it is uncommon on dry forest slopes or sometimes mesic forest at elevations of 180–730 m on Kaua‘i, O‘ahu, Moloka‘i, and Maui.

Metrosideros polymorpha Gaudich. ‘ōhi‘a, ‘ōhi‘a lehua

ENDEMIC | SECURE

Metrosideros—A genus of about 50 spe-

cies of trees, shrubs, and climbing lianas found in the Philippines, New Caledonia, New Zealand, and high volcanic islands of the Pacific. The name is from the Greek metra, heartwood, and sideron, iron, referring to the hard wood. There are generally considered to be 5 endemic species in the Hawaiian Islands, with one of them, M. polymorpha, exhibiting a wide ecological and morphological range of variation. Species concepts for the Hawaiian taxa are complex, and molecular studies suggest that the current taxonomy requires major revisions.

4.378 

Metrosideros polymorpha var. incana

Metrosideros macropus Hook. & Arn. ‘ōhi‘a, ‘ōhi‘a lehua

ENDEMIC | SECURE

to middle elevations on all of the major islands. The widespread var. glaberrima typically forms tall trees with glabrous leaves found at middle and upper elevations on all of the main islands except Ni‘ihau and Kaho‘olawe. Also widespread is the var. polymorpha, which forms shrubs to small trees with leaves typically densely pubescent on their lower surface on O‘ahu, Moloka‘i, Lāna‘i, Maui, and Hawai‘i. The dwarf var. pumila forms prostrate shrubs no more than 1 m tall with leaves either glabrous or pubescent on their lower surface, growing in bogs on Kaua‘i, Moloka‘i, and Maui. The var. dieteri comprises small trees with rough bark and unusually large glabrous leaves on Kaua‘i. The var. macrophyllum includes small to large trees with large glabrous leaves and bark separating into thin strips. It grows at middle elevations on Hawai‘i. The var. newellii forms shrubs to small trees with glabrous leaves growing along streams on Hawai‘i. Finally, var. pseudorugosa is a prostrate species with small leaves restricted to bogs on West Maui. The flowers of all forms secrete abundant nectar and are a favorite food source for many of the native honeycreepers. The flowers are widely used in lei making. The leaves were used as a remedy for thrush in children and also as an ingredient in a Hawaiian tonic to induce drowsiness. The wood has commercial use for paneling and floors. Ukulele keys are also made from the wood.

Metrosideros rugosa A. Gray lehua papa

ENDEMIC | SECURE

4.379 

4.377 

Metrosideos macropus

Small trees 5–7 m tall with fissured, flaky bark and persistent narrow scales surrounding the vegetative buds. Leaves ovate, 4.5–7.0 cm in length, glabrous. Flowers typically yellow but occasionally red. Found only in the Ko‘olau Mountains, O‘ahu.

Metrosideros polymorpha var. glaberrima

Tall trees, shrubs, or prostrate bog shrubs. Mature trees reach heights of 20–25 m and diameters up to 90 cm under optimal growing conditions. Leaves variable in size, shape, and pubescence. Flowers red, yellow, orange, or even white. Trees growing in wet soils often exhibit mats of aerial roots. Occurs in a wide variety of habitats at elevations from sea level to 2,200 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Morphologically and ecologically the most variable plant in the Hawaiian Islands. Eight varieties are recognized. The var. incana includes small trees or shrubs with leaves pubescent on their lower surface, and it occurs at low

4.380 

Metrosideros rugosa

FLOWERING PLANTS

Shrubs or sometimes even small trees with bark separating in thin flakes. Leaves obovate to suborbicular, 2–3 cm in length, under surface of the leaves densely woolly with tan to reddish brown hairs, veins strongly impressed on the upper surface and raised on the lower surface. Occurs near and on the summits of windward cliffs at elevations of 700–800 m in the Ko‘olau Mountains, O‘ahu.

Metrosideros tremuloides (A. Heller) Kunth. ‘āhihi, ‘āhihi kū ma kua, ‘āhihi lehua, ‘ōhi‘a ‘āhihi ENDEMIC | SECURE

length, glabrous. Occurs on high ridges at 600–1,300 m elevation on Kaua‘i, Moloka‘i, Lāna‘i, and West Maui. The var. waialealae is on Kaua‘i, while var. fauriei occurs on Moloka‘i, Lāna‘i, and West Maui.

Syzygium—A genus of trees and shrubs with about 1,200 species that are widely distributed in the paleotropics. Closely related to Eugenia. The name is from the Greek syn, together, and zygon, yoked, as the petals in some species are joined to form a cap. Many species are cultivated for their edible fruits. In the Hawaiian Islands, the rose apple (Syzygium jambos), the mountain apple (S. malaccense), and the Java plum (S. cumini) are naturalized. One species is endemic to the Islands.

Syzygium sandwicense (A. Gray) Mull. Berol. ‘ōhi‘a hā, hā, kauokahiki, pā‘ihi ENDEMIC | SECURE

4.381 

Metrosideros tremuloides

Shrubs to small trees. Leaves lanceolate, glabrous, often positioned at right angles to the stems. Occurs on steep slopes at elevations of 125–700 m in both the Ko‘olau and Wai‘anae mountains, O‘ahu. This species has a distinctive semiweeping habit, and the leaves often flutter in the wind like those of quaking aspen.

Nyctaginaceae

(FOUR-O’CLOCK FAMILY)

Herbs, shrubs, or trees. Key features are the often highly colored bracts and the petal-like calyx, the absence of petals, the herbaceous genera often scandent or sprawling, and the wood of the trees and shrubs usually soft and brittle. There are 27 genera and about 350 species found in tropical, subtropical, and temperate regions worldwide. Economically the family is of little importance except for a few genera that are cultivated, such as Bougainvillea, the four-o’clocks (Mirabilis), and the sand verbena (Abronia). In the Hawaiian Islands represented by 2 indigenous genera, Boerhavia and Pisonia. One species of Mirabilis has escaped and naturalized.

Boerhavia—A genus of about 50 species of erect to prostrate herbs, often with an enlarged taproot. The genus is named after Hermann Boerhaave, a seventeenth-century professor of botany at the University of Leiden. In the Hawaiian Islands, there are 2 indigenous, 1 endemic, and 1 introduced species. Many species are difficult to identify because of their polymorphic nature and the fact that they often intergrade with one another.

Boerhavia repens L. alena, nena

INDIGENOUS | SECURE

Metrosideros waialealae (Rock) Rock

4.383 

4.382 

Metrosideros waialealae var. waialealae

Shrubs to small trees with fissured and flaky bark. Leaves ovate to lanceolate or occasionally broadly ovate, 4–6 cm in

Syzygium sandwicense

Trees or shrubs 3–25 m tall with diameters up to 90 cm. Leaves coriaceous, sometimes concave, 2–14 cm in length, with a pink or red midrib. Occurs on ridges and slopes in mesic to wet forest and bogs at elevations of 230–1,220 m on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui. The wood was formerly used as fuel and in the construction of houses. A black dye for kapa (bark cloth) was made from the bark.

4.384 

Boerhavia repens

Robust perennial herbs from a thickened root, with many prostrate stems radiating from the root crown. Leaves ovate to suborbicular, 1–5 cm in length. Flowers in axillary cymes or pseudoumbels, pink to white. Native from Africa eastward to Polynesia and the Hawaiian Islands, where it occurs along shores and coastal areas

127

128

FLOWERING PLANTS

on Kure, Midway, and Pearl and Hermes atolls; on Lisianski, Laysan, French Frigate Shoals, and Nīhoa; and on all of the main islands. A highly variable and polymorphic species in the Hawaiian Islands.

Pisonia grandis R. Br. INDIGENOUS | SECURE

Pisonia umbellifera (G. Forst.) Seem. pāpala kēpau, pāpala

INDIGENOUS | SECURE

Pisonia—A genus of about 40 species of

shrubs, trees, and woody climbers found in the tropics, with a range centered in the neotropics but on a few Pacific islands. The name honors Willem Piso, a seventeenth-century Dutch physician, botanist, and pharmacist. In the Hawaiian Islands, there are 2 endemic species and 3 indigenous species that represent 3 separate colonizing lineages. The sticky fruits adhere easily to the feathers of birds for dispersal. The Native Hawaiians used the sticky gum of these seeds to capture native birds without harming them. Red and yellow feathers were highly prized for the feather cloaks of the ali‘i, and their collection was a specialized skill that was passed on through generations.

Pisonia brunoniana Endl. pāpala kēpau, pāpala

INDIGENOUS | SECURE

4.386 

Pisonia grandis

Large shrubs or trees up to 10 m or more in height. Wood very soft and brittle. Leaves typically opposite, sickle shaped, chartaceous to somewhat succulent in coastal areas, 9–30 cm in length, deciduous in very dry seasons. Flowers unisexual and plants dioecious or monoecious. Fruits very sticky and often seen adhering to the feathers of seabirds. Widely occurring from the western Indian Ocean to eastern Polynesia, particularly on small coral islands. Known from a single tree on Lisianski Island and historically from Maui.

Pisonia sandwicensis Hillebr.

āulu, kaulu, pāpala kēpau, pāpala ENDEMIC | SECURE

4.388 

Pisonia umbellifera

Large shrubs or small trees to 10 m tall, loosely branched. Leaves usually in false whorls of 3–8, sometimes opposite, variable in size and shape, 5–35 cm in length, 5–15 cm in width, thick. Plants monoecious. Fruit prism shaped and sticky. Occurs from New Hebrides, northern Queensland, Australia, Madagascar, and east to the Society and Hawaiian islands. In the Hawaiian Islands in wet to moist forest at elevations of 90–800 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Oleaceae

(OLIVE FAMILY)

4.385 

Pisonia brunoniana

Large shrubs or trees to 6 m tall, branchelets somewhat 4 angled. Leaves opposite or rarely in whorls of 3, broadly elliptic to obovate, up to 20 cm in length. Inflorescence a semi-open cyme, flowers perfect, moderately fragrant, white to greenish or bronze in color, becoming larger and more open when the plant is in fruit. Fruit very sticky. Native to Australia, New Zealand, the Kermadec, Norfolk, and Lord Howe islands, and the Hawaiian Islands. It occurs in dry to mesic areas at elevations of 425–1,220 m on O‘ahu, Moloka‘i, Lāna‘i, Maui, and Hawai‘i.

4.387 

Pisonia sandwicensis

Small trees 12–15 m tall, openly branched. leaves thick, coriaceous, oblong to broadly elliptic, 10–28 cm in length, petioles thick. Flowers unisexual, weakly fragrant. Fruit prism shaped and very sticky. Occurs in dry to mesic forest and rarely wet forest, usually on lower slopes or in gulches, at elevations of 260–1,220 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

A family of trees and shrubs or occasionally lianas. Key features include the typically 4-merous sepals and petals, the presence of 2 anthers, and opposite leaves. There are 24 genera and about 800 species found all over the world but particularly well represented in Asia and Malesia. The family is economically important for olives and olive oil (Olea), the lumber of ash (Fraxinus), and important ornamentals such as the lilac (Syringa), jasmine (Jasminum), golden bells (Forsythia), and Osmanthus. In the Hawaiian Islands represented by the indigenous genus Nes­ tegis. One species each of Fraxinus and Olea has naturalized.

Nestegis—A genus of 5 species of shrubs or trees on Norfolk Island, New Zealand, and the Hawaiian Islands. Closely related to Osmanthus. The name is from the Greek ne, not, and stegos, cover, as the type species lacks a corolla. In the Hawaiian Islands, there is 1 endemic species.

FLOWERING PLANTS

Nestegis sandwicensis (A. Gray) O. Deg., I. Deg. & L. A. S. Johnston olopua, pua, ulupua ENDEMIC | SECURE

a center of diversity in the Western Hemisphere. The genus was named for Christian Gottlieb Ludwig, a German botanist. In the Hawaiian Islands represented by Ludwigia octovalvis, which may have arrived in the Hawaiian Islands as an unintentional Polynesian introduction on taro plants. Seeds are buoyant but also stick easily to the feathers of migratory birds. It was first collected in the Hawaiian Islands in 1794.

Oxalidaceae

(WOOD SORREL FAMILY)

4.389 

Nestegis sandwicensis

Trees up to 8 m or more tall, rarely to 25 m, with thick, corrugated bark. Leaves coriaceous, lanceolate to narrowly elliptic, 7–22 cm in length, glossy on the upper surface. Flowers 7–11 in axillary inflorescences, 2–4 cm in length. Fruit turns from green to purplish black when mature. In dry to mesic forest on all of the main islands except Ni‘ihau and Kaho‘olawe. The dense wood was used for handles of adzes and other tools and as a rasp used in the manufacture of fishhooks. It is also excellent firewood, burning hot even when green.

Onagraceae

(EVENING PRIMROSE FAMILY)

A cosmopolitan family of annuals, perennial herbs, shrubs, and rarely trees, with a center of distribution in the temperate and subtropical Americas. There are 22 genera and about 650 species in the family. Many genera are cultivated, including woody Fuchsia and herbaceous Clarkia, Epilobium, Oenothera, and Ludwigia. In the Hawaiian Islands represented by the genus Ludwigia, which is either indigenous or an unintentional Polynesian introduction.

Ludwigia—A cosmopolitan genus of 82

species of generally aquatic plants, with many floating herbs and a few perennial species that are woody at the base. The genus has a pantropical distribution that extends to warm temperate regions, with

Herbaceous perennials or less commonly annuals or even woody plants, with a distribution in a range of tropical and subtropical habitats, particularly in the Southern Hemisphere. There are 5 genera and about 565 species. Leaves typically palmately compound and exhibiting nocturnal closure. Star fruit (Averrhoa) is a widely grown fruit, and tubers of Oxalis species were an important food source for indigenous Andean peoples. In the Hawaiian Islands represented by Oxalis corniculata, which may represent an unintentional Polynesian introduction or alternatively an indigenous species. It was collected in the Hawaiian Islands as early as 1779.

Papaveraceae (POPPY FAMILY)

Annual or perennial herbs, shrubs, or small trees. Key features are bisexual flowers with a calyx that falls away, the usually crumpled petals, numerous stamens, and a capsular fruit that dehisces by pores or flaps. Milky sap is often present. With an expanded concept of the family to now include the Fumariaceae, there are 43 genera and about 820 species distributed widely in temperate and tropical regions, primarily in the Northern Hemisphere. Economically the family is important for the production of opium and poppy seeds from Papaver somniferum and many ornamentals, such as oriental poppy (Papaver orientale), Iceland poppy (P. nudicale), bush poppy (Dendromecon), California poppy (Eschscholzia), blue poppy (Meconopsis), tulip poppy (Hunnemannia), and prickly poppy (Argemone). In the Hawaiian Islands represented by the indigenous genus Argemone and 3 introduced

genera that have naturalized: Bocconia, Eschscholzia, and Hunnemannia.

Argemone—A genus of about 23 species of

annual or perennial herbs with yellow to orange latex found in North and South America and the Hawaiian Islands. The name is from the Greek argema, cataract, as the sap of a poppylike plant was used to cure cataracts. The Hawaiian Islands have 1 endemic and 1 introduced species.

Argemone glauca (Nutt. ex Prain) Pope

kala, naule, pōkalakala, pua kala, prickly poppy ENDEMIC | SECURE

4.390 

Argemone glauca var. decipiens

Erect perennial herbs with bright yellow latex and prickly branches, stems somewhat woody at their base, 70–150 cm in length. Leaves conspicuously glaucous, covered with sharp, stiff prickles 8–21 cm in length. Petals 6, white. Two varieties are recognized and are differentiated by the number of prickles on the fruit. The var. decipiens is restricted to the leeward side of Hawai‘i and between Mauna Loa and Mauna Kea. The var. glauca occurs on all of the main islands in coastal dry forest, dry and subalpine woodland. The Native Hawaiians used the sap as a narcotic and analgesic for toothaches and ulcers and also as a remedy for warts.

Pentaphylacaceae

(PENTAPHYLAX FAMILY)

Small trees or shrubs. Key features are evergreen, entire leaves that are often unsymmetrical. Flowers generally perfect

129

130

FLOWERING PLANTS

but occasionally functionally dioecious; perianth parts imbricately arranged; anthers opening longitudinally. Pantropical in distribution and extending into temperate Asia. There are 12 genera and about 400 species, with a pantropical and warm temperate distribution. Economically the family has little importance but includes Cleyera, which is used as an ornamental. This family was formerly placed in a broader concept of the Theaceae. An older name for this segregate family is the Ternstoemiaceae. Represented in the Hawaiian Islands by the indigenous genus Eurya, which was placed in the Theaceae in the Manual. Eurya—A genus of evergreen trees and shrubs with about 70 species distributed in eastern Asia, Malesia, and the Pacific. Plants typically dioecious. The name is from the Greek euru, broad, referring to the wide sepals and petals of the plants. In the Hawaiian Islands represented by 1 uncommon endemic species.

Eurya sandwicensis A. Gray ‘ānini, wānini

ENDEMIC | RARE

veins reddish, lower surface conspicuously reticulate. Plants dioecious. Flowers 1–2 in leaf axils, nodding, petals 5, pale yellow to cream. Uncommon in mesic to wet forest and diverse mesic forest at elevations of 450–1,600 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i.

Phyllanthaceae

(PHYLLANTHUS FAMILY)

Trees, shrubs, herbaceous perennials, and occasionally climbers. Includes 53 genera and about 2,000 species, with more than half of the members in the genus Phyllanthus, which has as many as 1,200 species. The family was formerly included within the Euphorbiaceae. The group is distinguished by having 2 ovules in each locule of the ovary and an absence of milky sap. Most species are trees or woody shrubs, but some climbers and herbaceous taxa are present. Some species of Andrachne, Antidesma, Bischofia, Breynia, Margaritaria, and Phyllanthus are cultivated, and Antidesma, Baccaurea, Phyllanthus, and Uapaca provide edible fruit.

ha‘ā, ha‘āmaile, hame, mehame, mehamehame ENDEMIC | SECURE

Antidesma platyphyllum H. Mann

4.393 

ENDEMIC | SECURE

Trees 4–12 m tall with irregularly corrugated bark. Leaves thin, chartaceous, ovate to broadly elliptic, 7.5–15.0 cm in length, lower surface with tufts of hairs in the vein angles. Plants dioecious. Fruit fleshy, reddish to dark purple. Occasional in dry to mesic forest at elevations of 300–600 m or more on O‘ahu, Moloka‘i, Maui, and Hawai‘i.

Eurya sandwicensis

Small trees or shrubs, sometimes sprawling in windswept sites, stems 2–6 m in length. Leaves closely spaced, coriaceous, oblong to elliptic, 3.5–9.0 cm in length,

Antidesma pulvinatum Hillebr.

Antidesma—A genus of 170 species of dioecious trees of the paleotropics and subtropics, including the Pacific Basin. The name is from the Greek anti, against, and desma, headband. However, desma was meant to mean “poison” by a colleague of Linnaeus, supposedly because a member of the genus had been used as an antidote to snakebite. In the Hawaiian Islands, there are 2 endemic species representing a single colonizing lineage.

ha‘ā, ha‘āmaile, hame, mehame, mehamehame

4.391 

Trees 4–10 m tall. Leaves thick, ovate to obovate, glossy on the upper surface, 8–14 cm in length. Plants dioecious. Fruit fleshy, reddish to dark purple. Common in wet to mesic forest and in the Wahiawa Bog on Kaua‘i at elevations of 500–1,100 m. The var. hillebrandii is restricted to Kaua‘i, while the var. platyphyllum occurs on all of the main islands except Ni‘ihau and Kaho‘olawe. The Hawaiians used the logs as anvils for beating and preparing wauke fibers used to make kapa. A red dye was made from the fruit.

4.392 

Antidesma platyphyllum var. hillebrandii

Antidesma pulvinatum

Flueggea—A genus of 16 species of dioecious shrubs or trees ranging from the Mediterranean Basin to the paleotropics and Pacific Islands, with 1 endemic species in the Hawaiian Islands. Originally the Hawaiian plant now ascribed to this genus was described as the monotypic

FLOWERING PLANTS

Neowawraea by Joseph Rock and later erroneously transferred to Drypetes. The name honors Johann Flügge, a German physician and botanist.

qualities, or as ornamentals. The Hawaiian Islands have 1 endemic species and 2 species that have become naturalized weeds.

Flueggea neowawraea W. J. Hayden

Phyllanthus distichus Hook. & Arn.

ENDEMIC | ENDANGERED

ENDEMIC | SECURE

mēhamehame

pāmakani māhū

Phytolacca—A genus of about 25 species of perennial herbs, shrubs, or trees found in tropical and subtropical regions. The name is from the Greek phyton, plant, and the Latin lacca, the dye from lac insects, in reference to the staining juice of species in the genus. Phytolacca dioica is the famous ombu tree of Argentina, grown in the Hawaiian Islands as an ornamental. The Hawaiian Islands have 1 endemic species and 1 naturalized species.

Phytolacca sandwicensis Endl. pōpolo, pokeberry, pokeweed ENDEMIC | SECURE

4.394 

Flueggea neowawraea

Large trees up to 30 m tall and 2 m in diameter, bark rough and scaly, lenticels abundant. Leaves thin, chartaceous, ovate-elliptic, 4–14 cm in length. Plants dioecious. Fruit globose, reddish brown to black, juicy. Extremely rare in dry to mesic forest at elevations of 250–1,000 m on northwestern Kaua‘i, O‘ahu (Wai‘anae Mountains), Moloka‘i (extinct), East Maui (southwest slope of Haleakalā), and Hawai‘i (Kona Coast). One of the largest forest trees in the Hawaiian Islands, it is also one of the rarest, with only about 40 individuals remaining in the wild. Many of the older trees have died from insect attacks by the black twig borer (Xylosandrus compactus). Almost all of the remaining trees are basal sprouts from dead trunks or outplantings of saplings raised in cultivation.

Phyllanthus—A genus of about 1,200 species of herbs, shrubs, and trees mostly in the paleotropics but also represented in the neotropics and temperate North America and Pacific areas. The name comes from the Greek phyllon, leaf, and anthos, flower, because in some species the flowers are produced on the edge of leaflike branches. A few species are cultivated for their edible fruits, medicinal

4.395 

Phyllanthus distichus

Small shrubs 0.3–1.0 m tall. Leaves variable in shape and size, lanceolate to elliptic, 1.5–6.5 cm in length, distichous, deciduous. Usually monoecious but rarely dioecious. Fruit a small capsule. Occasional to common in mesic forest and occasionally dry shrubland at elevations of 60–950 m on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui.

Phytolaccaceae

(POKEWEED FAMILY)

Herbs, shrubs, trees, and lianas. Key features are the absence of petals, sepal-like perianths that are dry and inconspicuous or appear as petals, and a berrylike fruit often with colored juice and pulp. There are 16 genera and about 65 species found throughout the tropics and subtropics, particularly the neotropics. Economically the family is of no significant importance. A few species of Phytolacca, Rivina, Ercilla, and Agdestis are grown for ornament. Represented in the Hawaiian Islands by the indigenous genus Phytolacca and the naturalized Rivina humilis.

4.396 

Phytolacca sandwicensis

Perennial herbs 80–400 cm tall with sprawling stems. Leaves elliptic smooth or short-hairy, 8–22 cm in length. Flowers in racemes 8–22 cm in length, white to rose purple, fruit a dark purple berry. Occurs in open areas or streambeds in mesic to wet forest on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i. Often intergrading with the introduced southern pokeberry (P. octandra), making determinations difficult.

Piperaceae

(PEPPER FAMILY)

A tropical family of herbs, subshrubs, and shrubs, with a few species of lianas and trees. Key features are the often succulent and aromatic nature of the foliage, the densely and minutely spikelike inflorescence with naked flowers, and swollen nodes on the stems. There are 6 genera and about 2,150 species found in tropical regions around the world, particularly in wet shady locations. Economically the family is important for the condiment pepper (Piper nigrum), the

131

132

FLOWERING PLANTS

narcotic beverage kava, made from the roots of Piper methysticum, and a masticatory from the leaves of Piper betel. Several species of Peperomia are grown as house plants. In the Hawaiian Islands represented by the indigenous genus Peperomia and the Polynesian introduction of Piper methysticum.

Peperomia—A genus of succulent herbs that may be terrestrial or epiphytic. About 1,600 species are found throughout the tropics and subtropics of the world but with the greatest diversity in the neotropics. The name is from the Greek peperi, pepper, and homoios, resembling, because of the resemblance to the true pepper in the genus Piper. Represented in the Hawaiian Islands by 23 endemic, 2 indigenous, and 1 naturalized species. The native species are thought to derive from 4 separate colonizing ancestors.

Peperomia cookiana C. DC. ‘ala‘ala wai nui

ENDEMIC | SECURE

Peperomia globulanthera C. DC.

Peperomia hypoleuca Miq.

ENDEMIC | SECURE

ENDEMIC | SECURE

‘ala‘ala wai nui

4.398 

‘ala‘ala wai nui

Peperomia globulanthera

Succulent perennial herbs with erect to weakly ascending red stems 20–40 cm in length. Leaves 2–3(–6) per node, dark green above and pale green beneath, glabrous, 1.5–4.0 cm in length, palmately 3 veined. Spikes several, spreading, 3.5–6.0 cm in length. Epiphytic on trees or terrestrial in wet forests at elevations of 1,150– 2,070 m on Maui.

Peperomia hesperomannii Wawra ‘ala‘ala wai nui

ENDEMIC | SECURE

4.400 

Peperomia hypoleuca

Terrestrial or epiphytic succulent perennial herbs with erect to ascending red stems, 15–50 cm in length. Leaves 2–4 per node, 2–5 cm in length, dark green above and usually red with green veining below or sometimes entirely pale green. Terrestrial or epiphytic on trees in wet forest or occasionally mesic forests at elevations of 58–1,530 m on Hawai‘i.

Peperomia macraeana C. DC. ‘ala‘ala wai nui

ENDEMIC | SECURE

4.397 

Peperomia cookiana

Succulent perennial herbs with erect to weakly ascending red stems 10–35 cm in length. Leaves 2–4 per node, dark green above and pale green or partly red beneath, 1–5 cm in length, palmately 3 to 5 veined, typically pubescent. Spikes several to numerous, 3–9 cm in length. Terrestrial on wet rocks or epiphytic on trees in wet forest to mesic shrubland at elevations of 460–1,980 m on Kaua‘i, Moloka‘i, Maui, and Hawai‘i. A somewhat variable species with respect to leaf shape.

4.399 

Peperomia hesperomannii

Terrestrial or occasionally epiphytic succulent perennial herbs with reddish purple stems. Leaves whorled or opposite, 3–8 cm in length, yellowish green above and usually with some red beneath, palmately 3 nerved or 5 nerved in large leaves. Spikes 1 to several, 4–10 cm in length. A mostly terrestrial species from wet forests at elevations of 540–1,400 m on Kaua‘i.

4.401 

Peperomia macraeana

Terrestrial or rarely epiphytic succulent perennial herbs with tall, erect to ascending red stems, 50–200 cm in length. Leaves 2–4 per node, 5–15 cm in length,

FLOWERING PLANTS

thick and fleshy, green above, and green or with some red beneath, pinnately 5 nerved. Spikes several, 3–7 cm in length. Occurs in wet forests at elevations of 610–2,080 m on O‘ahu, Moloka‘i, Lāna‘i. Maui, and Hawai‘i. A variable species with respect to leaf shape and pubescence.

Peperomia membranacea Hook. & Arn. ala‘ala wai nui

ENDEMIC | SECURE

Succulent perennial herb with 1 to several ascending to erect stems up to 50 cm in length. Leaves whorled or opposite, dark green above and pale green below. Epiphytic or terrestrial, often growing on rotting logs in wet forest at elevations of 610–1,690 m on East Maui and in the Kohala Mountains, Hawai‘i.

Peperomia remyi C. DC. ‘ala‘ala wai nui

ENDEMIC | SECURE

Succulent perennial herbs with erect to ascending stems that are greenish, splotched with reddish purple. Leaves distinctive, narrowly linear-lanceolate, 12–20 cm in length. Rare and possibly now extinct, known only from mesic forest at elevations of 1,250–1,370 on East Maui (Kula Pipeline, lower Waikamoi). Sampled in the 1990s with a single population of less than 50 individuals. Recent surveys have found putative hybrids with Peperomia cookiana, but no individuals of the pure species.

Peperomia tetraphylla (G. Forster) Hook. & Ar n. ‘ala‘ala wai nui

INDIGENOUS | SECURE

4.402 

Peperomia membranacea

4.404 

Peperomia remyi

Succulent perennial herb with ascending to erect stems up to 80 cm in length. Leaves whorled on upper stem and opposite below, dark green on upper surface and pale green below. Epiphytic on trees and rocks or terrestrial in mesic to wet forest at elevations of 240–1,740 m on all of the main islands except Ni‘ihau, Lāna‘i, and Kaho‘olawe.

Succulent perennial herbs with erect to ascending reddish purple stems 20–50 cm in length. Leaves dark yellowish green on upper surface, pale below, veins often red, elliptic to obovate, 5–9 cm in length. Growing over rocks or terrestrial in mesic valleys and mesic to wet forest at elevations of 130–1,130 m on all of the main islands except Ni‘ihau, Lāna‘i, and Kaho‘olawe.

Peperomia obovatilimba C. DC.

Peperomia subpetiolata Yuncker

ENDEMIC | SECURE

ENDEMIC | EXTINCT?

‘ala‘ala wai nui

‘ala‘ala wai nui

4.406 

Peperomia tetraphylla

Succulent perennial herbs with several ascending or decumbent stems arising from a common base. Leaves usually in whorls of 4, either dark green or yellowish green above and green beneath. Pantropical in distribution and not closely related to other Hawaiian species. In the Hawaiian Islands in diverse habitats that range from shaded to open sites in mesic to wet forest, subalpine forest, and alpine desert at elevations of 70–2,290 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Pittosporaceae

(PITTOSPORUM FAMILY)

4.403 

Peperomia obovatilimba

4.405 

Peperomia subpetiolata

Trees, shrubs, or scandent subshrubs. Key features are the alternate coriaceous leaves crowded at branch tips and thus appearing to be whorled, 5 stamens with short filaments, and anthers opening by pores and a simple style. There are 7 genera and about 250 species distributed in paleotropical and warm temperate regions, particularly Australia. Economically the family is of little importance except for the cultivation of some species

133

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FLOWERING PLANTS

of Pittosporum and Sollya. In the Hawaiian Islands represented by the indigenous genus Pittosporum.

Pittosporum—A genus of about 200 species

of shrubs and trees found in the tropics and subtropics of Africa, Asia, Australia, New Zealand, and the Pacific Islands. The name is from the Greek pittos, pitch, and sporos, seeds, because the black seeds are covered with a sticky resin. Species are often distinguished by the size and shape of the capsule. In the Hawaiian Islands, there are 11 endemic species thought to derive from a single colonizing ancestor, and there are 2 naturalized species.

Shrubs or small trees 3–5 m tall. Leaves coriaceous, obovate, 9–30 cm in length, veins strongly impressed, glabrous above and reddish brown woolly below. Flowers in the leaf axils or along the stem, unisexual. In open bogs and wet forest at elevations of 640–1,520 m on Kaua‘i.

Pittosporum glabrum Hook. & Arn. hō‘awa, hā‘awa

ENDEMIC | SECURE

4.411 

Pittosporum confertiflorum A. Gray hō‘awa, hā‘awa

ENDEMIC | SECURE

4.409 

4.407 

Pittosporum confertiflorum

Shrubs or small trees 2–9 m tall. Leaves somewhat coriaceous, obovate, 7–30 cm in length, upper leaf surface with prominent impressed veins and reddish brown to brown matted wooly pubescence on both surfaces. The flowers are either terminal in the leaf axils or along the stem, perfect or unisexual. A widespread species in the Hawaiian Islands in dry to wet forest and subalpine forest at elevations of 180–2,200 m on O‘ahu, Lāna‘i, Maui, and Hawai‘i.

Pittosporum glabrum

Shrubs or small trees 4–8 m tall. Leaves chartaceous to semicoriaceous, upper surface with slightly impressed veins and nearly glabrous, lower surface glabrous in mature leaves but with a light woolly pubescence in young leaves. Flowers typically axillary, unisexual. The most widespread species in the Hawaiian Islands. In mesic to wet forest and margins of bogs at elevations of 240–1,970 m on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui.

Pittosporum hosmeri

Small trees 3–8 m tall. Leaves somewhat coriaceous, oblanceolate to obovate, 9–26 cm in length, smooth above and densely pale brown or whitish hairy beneath, margins typically inrolled. Flowers on stems or axillary, unisexual, and the plants dioecious. The capsules are the largest in the genus, 30–80 mm in length. Occurs in mesic to wet forest at elevations of 340– 1,070 m on Hawai‘i (Kohala Mountains south to Mauna‘anu, Ka‘ū Forest Reserve). The endemic Hawaiian crow, nearly extinct today, has been observed feeding on the oily seeds of this plant.

Pittosporum kauaiense Hillebr. hō‘awa, hā‘awa

ENDEMIC | SECURE

Pittosporum hosmeri Rock hō‘awa, hā‘awa

ENDEMIC | SECURE

Pittosporum gayanum Rock hō‘awa, hā‘awa

ENDEMIC | SECURE

4.412 

4.408 

Pittosporum gayanum

4.410 

Pittosporum hosmeri—fruit

Pittosporum kauaiense

Small trees 4–10 m tall. Leaves somewhat coriaceous, elliptic, 9–27 cm in length, veins moderately impressed above and with whitish to brownish matted hairs beneath. Occurs in diverse mesic forest and mesic valleys at elevations of 240–1,220 m on Kaua‘i. Often hybridizes with P. glabrum where the two are sympatric.

FLOWERING PLANTS

Pittosporum terminalioides Planch. ex A. Gray hō‘awa, hā‘awa

ENDEMIC | SECURE

as the source of psyllium from the seeds of Plantago psyllium used in laxative preparations. Many species have become cosmopolitan weeds. In the Hawaiian Islands represented by the genera Bacopa and Plantago.

Plantago hawaiensis (A. Gray) Pilg. laukāhi kuahiwi, plantain ENDEMIC | ENDANGERED

Bacopa—A mostly aquatic genus of 56

species widely ranging in tropical and subtropical areas but with a center of diversity in the Americas. Many species have alkaloids, saponins, and flavonoids with medicinal value influencing brain function. Some species are grown as aquarium plants, while others are considered to be noxious aquatic weeds. Bacopa is the Latinized form of an aboriginal name for these plants.

Bacopa monnieri (L.) Pennell ‘ae‘ae, water hyssop

INDIGENOUS | SECURE

4.413 

Pittosporum terminalioides

Trees 4–6 m tall. Leaves densely clustered at the ends of the branches, coriaceous and often thick, obovate to elliptic, 6.5– 15.0 cm in length, veins slightly impressed above, strongly so below, mature leaves glabrous above but with reddish brown to brown matted hairs below. Flowers axillary, unisexual. Occurs infrequently in dry to mesic forest and subalpine woodland at 300–2,010 m on Lāna‘i, Maui, and leeward Hawai‘i (Kīlauea to Kohala Mountains).

Plantaginaceae (PLANTAIN FAMILY)

Annual or perennial herbs, subshrubs (on islands), and a few aquatic species, with a broad cosmopolitan distribution. Key features are the usually rosulate foliage, with parallel venation, the inflorescences on stout or wiry stems, the flowers with 4 sepals and petals, and the stamens exserted on long, wiry filaments. Although once defined as a small family with only 3 genera, new molecular phylogenies include many genera here that were formerly placed in the Scrophulariaceae. Some systematists use the name Antirrhinaceae or Veronicaceae for this family. As newly circumscribed, there are 101 genera and about 1,900 species. Economically the family is important

4.415 

Plantago hawaiensis

Perennial herb arising from a stout woolly stem up to several cm in length. Leaves thick, basal, 7.5–22.0 cm in length. Flowers moderately to densely crowded. Similar in appearance to P. pachyphylla but present in much drier habitats. Occurs in dry to mesic shrublands, often in cracks of lava surfaces, at elevations of 1,800– 1,950 m on the leeward side of Hawai‘i. Five populations exist with about 5,000 plants, but major threats come from grazing by goats, mouflon sheep, and pigs.

Plantago pachyphylla A. Gray laukāhi kuahiwi, manene ENDEMIC | SECURE

4.414 

Bacopa monnieri

Perennial herbs with prostrate stems and fleshy leaves often forming mats. Widespread in tropical and subtropical regions. In the Hawaiian Islands in coastal areas on mudflats, rocks, marshes, and banks of brackish streams at elevations from sea level to 240 m on all of the main islands except Kaho‘olawe; also occurs on Midway Atoll. Popular as a garden groundcover. This genus was formerly placed in the Scrophulariaceae.

Plantago—A cosmopolitan genus of about 270 species of mostly annual or perennial herbs or rarely subshrubs. Plantago is from the Latin name for these plants. Represented in the Hawaiian Islands by 3 endemic species, 1 of them a subshrub and the other 2 stout perennial herbs. These are thought to derive from a single colonizing ancestor. There are 5 naturalized species.

4.416 

Plantago pachyphylla—habit

135

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FLOWERING PLANTS

Plumbago—A genus of perhaps 24 species of tropical and warm temperate perennials. The name is from the Latin plumbum, lead, and agere, to convey, in possible reference to the gray color imparted by the roots of these plants when handled. The latin agere can also be interpreted as meaning possessing the property of or resembling the color lead. In the Hawaiian Islands represented by 1 indigenous species. Plumbago auriculatum is cultivated.

Plumbago zeylanica L. 4.417 

‘īlie‘e

Plantago pachyphylla

Robust to small perennial herbs from a stout, unbranched woolly stem up to 20 cm in length. Leaves spirally arranged or 2 ranked, variable in shape, thick and coriaceous, oblong to lanceolate, 1.5–31.0 cm in length. Terrestrial or occasionally epiphytic on tree trunks, in wet forest, bogs, alpine shrubland, or alpine grassland at elevations of 700–2,200 m on Kaua‘i (Alaka‘i Swamp, Mount Wai‘ale‘ale), O‘ahu (Ko‘olau Mountains), Moloka‘i (Pēpē‘ōpae Bog), Maui, and Hawai‘i (Kohala Mountains, Kahuku Ranch). A highly variable plant whose many forms have been sometimes recognized at the species level.

Plantago princeps Cham. & Schltdl. ale, laukāhi kuahiwi

ENDEMIC | ENDANGERED

INDIGENOUS | SECURE

4.419 

Plantago princeps var. longibracteata

Small shrubs or robust perennial herbs with many-branched stems, 20–250 cm in length. Leaves tufted near the ends of the stems, thick, coriaceous, narrowly oblong-lanceolate to elliptic, 6–30 cm in length. Flowers densely crowded in spikes. Distinguished by its large branched stems. An uncommon species on steep slopes, rock walls, or at the bases of waterfalls at elevations of 480–1,100 m. Four varieties are recognized. The var. anomala is found on Kaua‘i (Hanapēpē and Kalalau valleys) and O‘ahu (Nāpepeiauolelo Gulch). The var. laxifolia occurs on Moloka‘i, Maui, and formerly on Hawai‘i. The var. longibracteata is present on Kaua‘i and O‘ahu, while the var. princeps occurs in both the Wai‘anae and Ko‘olau mountains of O‘ahu.

Plumbaginaceae (PLUMBAGO FAMILY)

4.418 

Plantago princeps var. anomala

Perennial herbs and shrubs, sometimes scandent. Key features are the presence of a 5-styled pistil, the leaves often with specialized chalk glands that excrete salts, and commonly with a long floral tube. There are 29 genera and about 730 species widely distributed but mostly of semiarid regions of the Old World. Economically the family is of little importance. Species of Plumbago and Limonium (statice of florists) are commonly cultivated. Represented in the Hawaiian Islands by the genus Plumbago.

4.420 

Plumbago zeylanica

Sprawling shrubs, stems 0.5–2.5 m in length. Leaves ovate, 3.0–12.5 cm in length, well developed on flowering stems. Flowers in racemes 6–30 cm in length, white with a sticky calyx. Widely distributed over the paleotropics and in the Hawaiian Islands occurring in arid, disturbed areas, sand dunes, dry forest, and shrubland at elevations from sea level to 610 m on all of the main islands. The sap was used to blacken tattoos.

Polygonaceae

(BUCKWHEAT FAMILY)

Annual or perennial herbs, shrubs, lianas, or rarely trees. Key features are the usual presence of stipules that are often united to form a sheath around the stem and a fruit that is usually a 3-sided achene or small nut. There are 46 genera and about 1,200 species that are cosmopolitan, with a distribution centered in north temperate areas. Economically the family is relatively unimportant. Buckwheat is obtained from Fagopyrum and rhubarb from Rheum. A few ornamentals such as Mexican creeper (Antigonon leptopus), the silver lace vine (Polygonum aubertii), and the sea grape tree (Coccoloba uvifera) are frequently cultivated. Many species

FLOWERING PLANTS

of Polygonum and Rumex are noxious weeds. Represented in the Hawaiian Islands by the indigenous genus Rumex. Species of Persicaria Rumex, Emex, and Polygonum have naturalized.

Rumex—A genus of annual or perennial herbs and rarely subshrubs, shrubs, or lianas, with about 200 species that are worldwide in distribution but with the largest number of species in temperate regions. Rumex is the old Latin name for these plants. Many species are cosmopolitan weeds. In the Hawaiian Islands, there are 3 endemic species representing a single colonizing lineage and 5 naturalized species.

Scandent shrubs or lianas with long stems usually supported by trees in forested areas or suberect at high elevations. Leaves oblong-ovate to deltate, 6–22 cm in length. Flowers unisexual, and both sexes occur on the same plant in large, usually red spreading inflorescences. Occurs in mesic to wet forest and subalpine woodland at elevations of 660–3,050 m on Moloka‘i, Maui, and Hawai‘i. A variable and uncommon plant.

Rumex skottsbergii O. Deg. & I. Deg. pāwale, dock, sorrel ENDEMIC | SECURE

petals, and often numerous stamens. Traditionally the family has included 20 genera and about 325 species that are cosmopolitan in distribution but particularly well represented in western North America and in the Andes of South America. Recently most genera were moved to the Montiaceae, Didiereaceae, Anacampserotaceae, and Talinaceae, thus making the family monotypic and containing only the genus Portulaca. Herbage of the pigweed (Portulaca oleracea) is used as a salad green. In the Hawaiian Islands represented by the indigenous genus Portulaca.

Portulaca—A genus of annual or perennial herbs or sometimes low shrubs, with 107 species found throughout the tropics and warm temperate regions. Portulaca is the Latin name for pigweed or purslane (P. oleracea), which is sometimes used as a salad. There are 3 endemic, 1 indigenous, and 3 naturalized species in the Hawaiian Islands. The native species represent 2 separate colonizing lineages.

Rumex albescens Hillebr. hu‘ahu‘akō, dock, sorrel ENDEMIC | SECURE

Portulaca lutea Sol. ex G. Forst. ‘ihi

INDIGENOUS | SECURE

4.421 

Rumex albescens

Subshrubs with decumbent stems up to 2 m in length, glabrous to moderately pubescent. Leaves lanceolate and up to 30 cm or more in length but variable in shape and leaf margin, which varies from entire to undulate or crisped. Scattered in mesic forest and in dry shrubland and cliff faces on Nīhoa, Kaua‘i, and the Wai‘anae Mountains, O‘ahu.

Rumex giganteus W. T. Aiton

pāwale, uhauhakō, dock, sorrel ENDEMIC | SECURE

4.423 

Rumex skottsbergii

Shrubs with stiffly erect stems 70–100 cm in length. Leaves thick, ovate to elliptic, 5.5–9.0 cm in length. Flowers greenish in erect, compact inflorescences. Occurs on dry lava at elevations of 460–1,300 m on Hawai‘i (North and South Kona, Puna, and southern Ka‘ū districts). Similar to the preceding species, especially where R. giganteus grows on Mauna Kea and Haleakalā, but differing in its more compact greenish inflorescence.

Portulacaceae

(PURSLANE FAMILY)

4.422 

Rumex giganteus

Annual or perennial herbs and occasionally subshrubs. Key features are the usual presence of succulent leaves, 2 sepals, 5

4.424 

Portulaca lutea

Prostrate or somewhat ascending perennial herbs with succulent stems having a thin, pale, corky layer on older parts. Leaves more or less uniformly spaced on the stems. Flowers yellow. Widespread in the Pacific. Occurs in coastal and strand habitats on lava, raised coral reefs, and sand dunes at elevations from sea level to 40 m on all of the Northwestern Hawaiian Islands except Pearl and Hermes Atoll and on all of the main islands except Kaho‘olawe.

137

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FLOWERING PLANTS

Portulaca molokiniensis Hobdy

Portulaca villosa Cham.

ENDEMIC | RARE

ENDEMIC | VULNERABLE

‘ihi

‘ihi

Embelia—A genus of about 130 species

of scandent shrubs or lianas in the paleotropics and Pacific Islands. This genus was formerly placed in the Myrsinaceae, a family now included in the Primulaceae. The name is from the Greek em, within, and belos, dart, referring to the secretory lines on the leaves. In the Hawaiian Islands known only from 1 endemic species.

Embelia pacifica Hillebr. kilioe

ENDEMIC | SECURE

4.425 

Portulaca molokiniensis

Stout perennial herbs with stems branched at or beneath the ground, often forming clumps. Leaves clustered at the ends of the branches. Flowers in dense headlike clusters, yellow. Rare in coastal sites at 10–115 m elevation on steep rocky slopes, at the base of sea cliffs, and other coastal habitats on Molokini, Pu‘ukoa‘e Islet, and Kamōhio Bay, Kaho‘olawe.

Portulaca sclerocarpa A. Gray ‘ihi, ‘ihi mākole, po‘e

ENDEMIC | ENDANGERED

4.427 

Portulaca villosa

Perennial herbs from a fleshy, tuberous taproot with prostrate to ascending stems and succulent, linear leaves. Flowers white, pink, or pink with a white base. Occurs on dry rocky clay, lava, or coralline reef sites at elevations from sea level to 300 m on Nīhoa, Ka‘ula (islet southwest of Ni‘ihau), and on all of the main islands except Ni‘ihau and Kaho‘olawe.

Primulaceae

(PRIMROSE FAMILY)

4.426 

Portulaca sclerocarpa

Perennial herbs from a fleshy, tuberous taproot with prostrate to ascending stems and succulent, linear leaves. Flowers white, pink, or pink with a white base. Uncommon in dry habitats such as subalpine woodland, bare cinders, and volcanic steam vents at elevations of (30) 1,030–1,630 m on Lāna‘i (Po‘opo‘o Islet) and Hawai‘i. Closely related to P. villosa but differing in its thick-walled, indehiscent capsules. There are 10 known populations that total somewhat more than 1,000 plants. Threats come from feral pigs, goats, mouflon sheep, and fire.

Herbs and subshrubs; widely distributed. Key features are the presence of secretory cells that contain red substances in many genera, a single whorl of stamens opposite the corolla lobes, and a dehiscent capsular fruit containing numerous seeds. Based on new molecular data, the family has recently been expanded to include the Myrsinaceae and Theophrastaceae. With these additions, there are 60 genera and about 2,575 species worldwide but primarily distributed in tropical and north temperate regions. Economically the family is important for timber (Myrsine) and for ornamentals in genera such as Primula, Cyclamen, Anagallis, and Lysimachia. In the Hawaiian Islands represented by the indigenous genera Embelia, Lysimachia, and Myrsine and the naturalized weedy Anagallis arvensis.

4.428 

Embelia pacifica

Lianas or scandent shrubs with young branches covered with prominent lenticels. Leaves membranous, elliptic to obovate, 3.5–19.0 cm in length, conspicuously glandular-punctate. Flowers in axillary racemose inflorescences, 15–45 cm in length. Fruit a drupe turning dark purple when mature. Occasional in mesic or sometimes wet forest at elevations of 450–1,650 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Lysimachia—A genus of about 190 species of perennial herbs, subshrubs, or shrubs that are cosmopolitan in distribution but with the Himalayas of southwest China being the center of species diversity. Leaves may be alternate, opposite, or whorled. Flowers in terminal racemes, corymbs, panicles, or solitary in the leaf axils, variable in color. Special flower glands called elaiophores are particularly attractive to bees. The name honors Lysimachus, the king of Thrace, who reportedly waved the plant in front of a bull to pacify the animal. In the Hawaiian Islands represented by 13 endemic and 1 indigenous species. The endemic species represent a single colonizing lineage distinct from the indigenous species. Only 11 species were listed in the Manual.

FLOWERING PLANTS

Lysimachia glutinosa Rock

Lysimachia kalalauensis Skottsb.

ENDEMIC | SECURE

ENDEMIC | SECURE

from Mauritius, Reunion, eastern Africa, China, Japan, Taiwan, to Pacific Islands. In the Hawaiian Islands rare on coastal sites on Ni‘ihau, Kaua‘i, Moloka‘i, Maui, and northwest Hawai‘i. This species represents an independent colonizing ancestor from the other Hawaiian species.

Lysimachia pendens K. L. Marr ENDEMIC | RARE

4.431 

4.429 

Lysimachia glutinosa

Erect to sprawling shrubs 0.6–2.0 m tall with viscid stems, bark reddish brown. Leaves alternate, well spaced, oblanceolate to elliptic, 6–15 cm in length. Flowers solitary in the leaf axils, sweetly scented, corolla cream colored. Occurs in diverse mesic forest at elevations of 1,090–1,220 m in the Kōke‘e area of Kaua‘i.

Lysimachia kalalauensis

Sprawling to scandent shrubs with fewbranched stems 2–3 m in length. Leaves alternate, well spaced, thin, coriaceous, elliptic, 5.0–9.5 cm in length. Flowers solitary in the leaf axils, corolla green with purple veining, pedicel elongating in fruit. Rare in shaded sites in diverse mesic forest on Kaua‘i (Kōke‘e, from Kalalau to Mākaha valleys, Hanapēpē Valley).

Lysimachia mauritiana Lam. INDIGENOUS | SECURE

Lysimachia hillebrandii J. D. Hook. ex A. Gray kolekole lehua, kolokolo kuahiwi ENDEMIC | SECURE

4.433 

Lysimachia pendens

Small matlike shrubs with delicate pendulous or prostrate branches up to 60 cm long. Differs from L. filifolia in having narrowly lanceolate and tomentose leaves 2–4 mm wide compared to filiform leaves less than 2 mm wide in the latter. Known only from several small populations at 720 m elevation at the headwaters of the north fork of the Wailua River, Kaua‘i. This species was first discovered in 1987 and not included in the Manual.

Lysimachia remyi Hillebr. ENDEMIC | SECURE

4.430 

Lysimachia hillebrandii

Sprawling shrubs with reddish brown bark and many-branched stems, stems 0.4–2.5 m in length. Leaves alternate, closely to well spaced, ovate to elliptic, 2.5–9.0 cm in length. Flowers solitary in the leaf axils, corolla reddish purple to burgundy. Occurs in open sites in mesic to wet forest at elevations of 610–1,500 m on Kaua‘i (Hā‘upu to Limahuli), O‘ahu, eastern Moloka‘i, Lāna‘i, and Maui. Closely related to L. remyi, which tends to have narrower, more closely spaced leaves.

4.432 

Lysimachia mauritiana

Somewhat fleshy perennial herbs becoming woody at the base, stems 10–80 cm in length. Leaves alternate, chartaceous to somewhat fleshy, spathulate to elliptic, 2–6 cm in length, lower surface dotted with black glands. Flowers in congested racemes, corolla white or tinged pink, calyx dotted with black glands. Native over a wide geographic range

4.434 

Lysimachia remyi

139

140

FLOWERING PLANTS

Erect, many-branched shrubs, 50–200 cm tall, sometimes sprawling with a few basal branches, young shoots with rust-colored pubescence. Leaves typically imbricate and closely spaced, linear-filiform to narrowly elliptic, 1.5–5.5 cm in length. Flowers solitary in the leaf axils, corolla reddish purple. Scattered in open sites, often on wet cliffs or rock walls in mesic shrubland at elevations of 760– 2,200 m on Moloka‘i and Maui.

Many-branched shrubs or small trees 2–6 m tall. Leaves narrowly elliptic to oblanceolate, 1.5–4.2 cm in length, yellowish green, well distributed along the branches. Flowers apparently perfect. Occurs in mesic forest or sometimes in wet forest at elevations of 580–1,220 m in the Kōke‘e and Hā‘upu Ridge area of Kaua‘i.

Myrsine—A genus of about 200 species

ENDEMIC | SECURE

of trees and shrubs in tropical and subtropical areas and throughout the Pacific Basin. Leaves simple and alternate, punctuate, and with secretory canals. Flowers perfect or unisexual, with the plants then dioecious. Flowers of many Hawaiian species have pronounced secretory lines along the petals, which aid in pollination. Fruit is a semiglobose drupe. This genus was formerly placed in the Myrsinaceae, but the family is now included within the Primulaceae. The name is from a Greek name for myrtle, not applicable to these plants. In many species, the flowers are cauliflorous. Species differences are in vegetative traits and often subtle. In the Hawaiian Islands represented by 19 closely related species representing a single colonizing lineage. There is a rich assemblage of 14 species on Kaua‘i. Thirteen species are single-island endemics.

Myrsine alyxifolia Hosaka kōlea

ENDEMIC | SECURE

Myrsine helleri (O. Deg. & I. Deg.) H. St. John kōlea, ‘ōliko

Small trees 3–6 m tall with branches that have conspicuous lenticels. Leaves bright red when young or at least with a purple midrib, which is pubescent below, thinly fleshy, elliptic to oblong-ovate, oblanceolate to narrowly elliptic, 8–16 cm in length. Flowers apparently perfect. Occurs in dry forests to occasionally mesic forests, often on open lava fields at elevations of 300–1,000 m on the leeward sides of all of the main islands except Ni‘ihau and Kaho‘olawe.

Myrsine lessertiana A. DC. kōlea, kōlea lau nui ENDEMIC | SECURE

4.436 

Myrsine helleri

Branched shrubs 2–5 m tall with dark reddish brown branches. Leaves narrowly lanceolate to linear-lanceolate, 4–7 cm in length, with pronounced black glandular dots on the upper surface. Flowers apparently perfect. Occurs in and on the margins of the Wahiawa Bog at an elevation of 640–650 m and the area of the Alaka‘i Swamp to Mount Wai‘ale‘ale on Kaua‘i at 1,400–1,525 m.

4.438 

Myrsine lessertiana—habit

4.439 

Myrsine lessertiana

Myrsine lanaiensis Hillebr. kōlea

ENDEMIC | SECURE

4.435 

Myrsine alyxifolia

4.437 

Myrsine lanaiensis

FLOWERING PLANTS

Trees or sometimes shrubs 2–8 m tall. Leaves dark green with young leaves reddish or reddish purple, clustered at the tips of the branches, coriaceous, oblanceolate to oblong-ovate, 5–12 cm in length. Flowers apparently perfect or questionably unisexual. Occurs primarily in mesic to wet forests but sometimes in dry forests to subalpine woodlands at elevations of 300–2,290 m on all of the main islands except Ni‘ihau and Kaho‘olawe. A highly variable plant throughout its range. The wood was formerly used for house posts and beams and for anvils to beat kapa. A red dye for kapa was made from the sap and a black kapa dye was made from the charcoal.

Myrsine punctata (H. Lév.) Wilbur ENDEMIC | SECURE

Occurs in mesic to wet forest and bogs at elevations of 200–1,470 m on O‘ahu, Moloka‘i, Lāna‘i, Maui, and Hawai‘i.

Myrsine wawraea (Mez) Hosaka ENDEMIC | SECURE

Myrsine linearifolia Hosaka ENDEMIC | ENDANGERED

4.441 

Myrsine punctata

Many-branched shrubs 1–3 m tall, with branches brown to reddish brown. Leaves closely spaced on upper branches, slightly fleshy, oblanceolate to linear-oblanceolate or elliptic, 0.6–2.5 cm in length. Flowers apparently perfect. Occurs in mesic to wet forest and along the margins of bogs at elevations of 1,160–1,280 m on Kaua‘i (Kōke‘e Plateau, Alaka‘i Swamp, Mount Kāhili) and the Wai‘anae Mountains (Kuaokalā), O‘ahu.

Myrsine sandwicensis A. DC. kōlea, kōlea lau li‘i

ENDEMIC | SECURE

4.440 

Myrsine linearifolia

Branched shrubs 3–8 m tall with ascending green to reddish brown branches. Leaves well distributed on upper branches, slightly fleshy, linear, 5–9 cm in length, yellowish purple near the base. An uncommon species in diverse mesic forest to wet forest at elevations of 580– 1,280 m on Kaua‘i (west half to Pu‘ukila). Recent censuses report 12 existing populations with a total of less than 200 individuals. Threats come from feral pigs and alien plant species.

4.442 

Myrsine sandwicensis

Many-branched shrubs or small trees 1.5–4.0 m tall. Leaves well distributed along the branches, dark green, coriaceous, oblanceolate to obovate, 1–3 cm in length. Flowers apparently perfect.

4.443 

Myrsine wawraea

Sprawling shrubs or small trees typically only 0.3–1.0 m tall, branches few, green to reddish brown. Leaves dark green, appearing whorled because of short internodes, semicoriaceous, oblanceolate to elliptic, midrib typically dark purple, 7–18 cm in length. Flowers unisexual and the plants thus dioecious. Occurs in mesic to wet forest and along the margin of bogs at elevations of 1,020–1,400 m on Kaua‘i (Kōke‘e, Alaka‘i Swamp, Mount Kāhili, Kalalau Trail).

Ranunculaceae

(BUTTERCUP FAMILY)

Herbs, both aquatic and terrestrial, occasionally vines and shrubs. Key features are the typically divided or compound nature of the foliage, the mostly numerous, spirally arranged stamens, and a gynoecium of several to many pistils. There are 56 genera and about 2,100 species worldwide, primarily in temperate, boreal regions and tropical mountains. Economically the family is important for the large number of ornamentals, such as buttercups (Ranunculus), windflower (Anemone), larkspur (Delphinium), meadow rue (Thalictrum), columbine (Aquilegia), and virgin’s bower (Clematis). Alkaloids are widespread in the family, as in goldenseal (Hydrastis) and wolfbane (Aconitum), and many have found use in traditional

141

142

FLOWERING PLANTS

medicine. In the Hawaiian Islands represented by the indigenous genus Ranunculus and naturalized Anemone.

Ranunculus—A large genus of about 600

species of terrestrial or aquatic herbs centered in distribution in north temperate and boreal habitats but also tropical mountains. The name is from the Latin rana, frog, because many species are found in wet or aquatic environments. In the Hawaiian Islands, there are 2 rare endemic species representing a single colonizing lineage, and there are 4 naturalized species.

Ranunculus hawaiensis A. Gray ENDEMIC | ENDANGERED

often possessing a hypanthium on which the calyx, corolla, and stamens arise, and the stamens opposite the petals, which are often strongly concave. There are 53 genera and about 875 species distributed worldwide but more common in tropical and subtropical regions. Economically the family is important for ornamental shrubs such as buckthorns (Rhamnus), Ceanothus, jujube (Zizyphus), and the raisin tree (Hovenia). A laxative, cascara (sagrada), is obtained from Rhamnus purshiana. In the Hawaiian Islands represented by the indigenous genera Alphitonia, Colubrina, and Gouania and the introduced Rhamnus californica.

United States, the Caribbean, and South America. Other species occur in Malesia, Africa, and islands of the Pacific and Indian oceans. The name is from the Latin coluber, snake, although its application to these plants is not known. In the Hawaiian Islands represented by 1 rare endemic and 1 indigenous species, each representing an independent colonizing lineage.

Colubrina asiatica (L.) Brongn.

‘ānapanapa, kauila ‘ānapanapa, kauila kukuku INDIGENOUS | SECURE

Alphitonia—A genus of 15 species of trees

and shrubs in Malesia, eastern Australia, and the Pacific Islands. The name is from the Greek alphiton, barley meal, because the mesocarp of these plants is dry and mealy. In the Hawaiian Islands represented by 1 endemic species.

Alphitonia ponderosa Hillebr. kauila, o‘a

ENDEMIC | RARE

4.444 

Ranunculus hawaiensis

Erect to ascending perennial herb with fibrous roots and basal leaves twice ternately compound. The flowers are small in loose, branched inflorescences. A rare species in open grassy or loose rocky slopes in mesic to wet forest at elevations of 1,820–2,040 m on East Maui and Hawai‘i. The most recent survey failed to find plants at two former locations on Maui and reported only 14 individuals fragmented in 6 populations on Hawai‘i. Threats have come from feral animals, slugs, and alien plant species.

Rhamnaceae

(BUCKTHORN FAMILY)

Trees, shrubs, subshrubs, or even perennial herbs. Some genera climb by way of coiled tendrils or hooks. Key features are the simple, entire leaves, the flowers

4.445 

Alphitonia ponderosa

Trees 4–25 m tall with whitish, corrugated bark. Leaves somewhat coriaceous, ovate to lanceolate, 8–15 cm in length, lower surface with dense rusty tomentum on young leaves. Flowers in an elongate paniculate inflorescence, 6–8 cm in length. The hard, black, round fruit contains seeds covered by a red aril. Occurs in dry to mesic forest at elevations of 240–1,250 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Moderately common on Kaua‘i but rare on the other islands. The hard wood was formerly used for house beams, kapa mallets, javelins, and digging sticks.

Colubrina—A genus of 33 species of trees or shrubs primarily from the western

4.446 

Colubrina asiatica

Weakly climbing to scandent shrubs, young branches zig-zag. Leaves alternate, thin, glossy, ovate, 3.5–11.5 cm in length, margins serrate. Flowers usually perfect. Fruit globose, explosively dehiscent. Native in coastal and lowland habitats in Africa, Indian Ocean islands, Malesia, Australia, and the Pacific Islands including the Hawaiian Islands. In strand and coastal sites at elevations from sea level to 100 m on Ni‘ihau, Kaua‘i, O‘ahu, Lāna‘i, Moloka‘i, and West Maui (Launiupoko). The toxins in the plant were formerly used by the Hawaiians to anesthetize fish to allow their capture.

Colubrina oppositifolia Brongn. ex H. Mann kauila

ENDEMIC | ENDANGERED

4.447 

Colubrina oppositifolia

FLOWERING PLANTS

Trees 3–13 m tall. Leaves decussate, dull, ovate, 6–12 cm in length, glandular beneath. The broad, rounded fruit explodes when ready to open. Rare in dry to mesic forest at elevations of 240–920 m in the Wai‘anae Mountains, O‘ahu, West Maui (Honokōwai), and leeward Hawai‘i (see figure 8.4). This tree has very hard wood and took the place of metal implements in ancient Hawaiian culture. It was used to make kapa beaters, spears, and other weapons and for poles for kāhili, the symbol of royalty. Once a common dry forest tree, native populations have sharply declined; recent surveys have reported only 10 populations with a total of less than 300 individuals.

Gouania—A pantropical genus of about 50 species of shrubs or lianas. Saponins, soapy compounds, are present in the bark and used in some regions for shampoo or cleaning teeth. The name honors Antoine Gouan, an eighteenth-century professor at Montpellier, France. In the Hawaiian Islands represented by 2 endemic shrubs and 1 endemic liana with coiled tendrils, all rare. These species represent 2 separate colonizing lineages.

Gouania hillebrandii Oliver ENDEMIC | ENDANGERED

Waiakuilani Gulch), Lāna‘i (now extinct), West Maui (Pa‘upa‘u and Līhau Mountain, Lahaina District), Kaho‘olawe (now extinct), and Hawai‘i. Surveys a decade ago found only 3 extant populations with an estimated 1,700–2,100 individuals, all in the Lahaina District of West Maui.

Gouania meyenii Steud. ENDEMIC | ENDANGERED

Rosaceae

(ROSE FAMILY)

4.449 

Gouania meyenii

Erect to spreading shrubs, stems up to 2 m or more in length. Leaves chartaceous, ovate to rounded, blades 3.0–7.5 cm. Tendrils absent. Fruit 2 winged. Very rare and known only from a few populations in mesic forests at an elevation of about 800 m in the Hanalei and Waimea districts of Kaua‘i and the Wai‘anae Mountains, O‘ahu. Recent estimates suggest a total population of 38–63 individuals split between the two islands. Threats have come from feral pigs and goats, alien plant species, and fire.

Gouania vitifolia A. Gray ENDEMIC | ENDANGERED

4.448 

Gouania hillebrandii

Erect to sprawling shrub forming clumps, branches up to 2 m in length. Leaves, elliptic, chartaceous to coriaceous, 2–6 cm in length. Flowering branches without tendrils. Fruit 3 winged. Very rare and known from only a few populations, but it was once more widespread. Occurs in dry forest on Moloka‘i (Kamalō,

form dense clumps. Leaves chartaceous, broadly ovate, blades 3–8 cm in length, underside with dense mat of soft hairs. Flowers borne on short lateral branches with 1–2 leaves and stiff, recurved tendrils. Known only from a few collections, but it once occurred on O‘ahu (Wai‘anae Mountains), West Maui (Lahaina), and Hawai‘i (Ka‘ū). Thought to be extinct until rediscovered in the Wai‘anae Mountains and on Hawai‘i in disturbed mesic forest. Detailed census data is difficult because of the vegetative reproduction of the plants, but no more than 100 individuals survive. Threats come from feral pigs and goats, alien plant species, and potentially fire. This species is considered to represent an independent colonizing lineage from the other 2 species.

Trees, shrubs, or herbs. Key features are the typical presence of stipules, a hypanthium in most of the genera, and flowers with 5 petals and sepals. The family has sometimes been divided into several families, depending on the authors. As recognized here, there are 85 genera and about 3,000 species that are found throughout the world, but its greatest diversity is in temperate and subtropical areas of the Northern Hemisphere. Economically the family is very important for a number of edible fruits, such as apples (Malus), pears (Pyrus), cherries, plums, apricots, almonds, and prunes (Prunus), loquats (Eriobotrya), strawberries (Fragaria), and blackberries and raspberries (Rubus). The family includes a number of ornamentals, such as fireberry (Pyracantha), hawthorn (Crateagus), flowering quince (Chaenomeles), rose (Rosa), and spiraea (Spiraea). In the Hawaiian Islands represented by 4 indigenous genera: Acaena, Fragaria, Osteomeles, and Rubus. One species of Pyracantha has naturalized.

Acaena—A genus of perennial herbs,

4.450 

Gouania vitifolia

Climbing shrubs with ridged stems, sometimes vegetatively reproducing to

sometimes woody at the base. There are about 100 species mostly in the Southern Hemisphere, with 1 species in the Hawaiian Islands and 1 in California. The name is from the Greek akiana, thorn, referring to the spiny nature of the receptacle. In the Hawaiian Islands represented by 1 endemic species.

143

144

FLOWERING PLANTS

Acaena exigua A. Gray liliwai, nani Wai‘ale‘ale ENDEMIC | EXTINCT?

Small tufted perennial with densely leafy stems and pinnately compound leaves with a glossy upper surface. Often growing with mosses and liverworts. Originally collected in bogs on Kaua‘i (Mount Wai‘ale‘ale), where it was last seen in 1870 and West Maui (Pu‘ukukui), where it was last collected in 1957 until a single plant was rediscovered in 1997. Despite the presence of appropriate bog habitat on both Kaua‘i and West Maui, no new observations of this small plant have been made, and it is best considered extinct. Not pictured.

Fragaria—A genus of perennial herbs arising from scaly rhizomatous rootstocks and often propagating by means of stolons. There are about 20 species and many cultivars found in Eurasia, North America, the Andes of Chile and Argentina, and 1 indigenous species in the Hawaiian Islands. The name is from the Latin fragrum, fragrant, because of the sweet smell of the fruit. In addition to the native species, the European strawberry has naturalized on Kaua‘i and Hawai‘i. Strawberries are also cultivated commercially in the Islands.

Fragaria chiloensis (L.) Duchesne subsp. sandwicensis (Decne.) Staudt. ‘ōhelo papa, strawberry

INDIGENOUS | VULNERABLE

4.451 

Fragaria chiloensis

Perennial herbs from a short rootstock that also produces long stolons. The red fruit is edible. Native to the Pacific Coast of North America from Chile to Argentina, the Juan Fernandez Islands, and the Hawaiian Islands. In dry disturbed areas, wet forest, and subalpine shrub­ land at elevations of 1,160–3,070 m on East Maui and Hawai‘i. Probably one of

the parents of the strawberries grown for food. Hawaiian plants are the endemic subsp. sandwicensis. Very similar plants occur on the Juan Fernandez Islands.

Rubus hawaiensis A. Gray ‘ākala, ‘ākalakala

ENDEMIC | SECURE

Osteomeles—A genus of 3 species of ever-

green or deciduous shrubs of Eastern Asia and Polynesia, but sometimes interpreted as a single highly variable species. The name is from the Greek osteon, bone, and melon, apple, referring to the nature of the fruit. In the Hawaiian Islands, there is 1 widespread indigenous species.

Osteomeles anthyllidifolia (Sm.) Lindl. ‘ūlei

INDIGENOUS | SECURE

4.453 

4.452 

Osteomeles anthyllidifolia

Shrubs with prostrate stems but occasionally becoming erect. The dark green pinnate leaves are coriaceous and glossy above. The fruit is white at maturity with white or purple flesh. Occurs in the Cook Islands, Tonga, and in the Hawaiian Islands, where it is found in a wide variety of habitats from along the coast to subalpine shrubland at elevations of from 2 to 2,320 m on all of the main islands except Ni‘ihau and Kaho‘olawe. The fruit is insipid but edible, and it was also strung into lei. The wood was made into digging sticks, fish spears, and other implements, while the slender branches were bent into hoops for fishnets.

Rubus—A cosmopolitan genus of erect, scandent, or prostrate shrubs with prickly stems, except in the Hawaiian species R. hawaienesis. There may be as many as 250 species, but because of hybridization and apomixis it is difficult to apply the species concept in the strict sense of the word. The name is from the Latin meaning bramble. In the Hawaiian Islands, there are 2 endemic species that represent independent colonization events. There are 5 naturalized species, 2 of which have become serious pests in the Islands.

Rubus hawaiensis

Shrubs with shredding bark and erect to arching stems. The palmately compound leaves are generally without prickles. The flowers are usually dark pink to rose or sometimes white; fruit is dark red to dark purple or yellow and edible, although rather bitter. Occurs in mesic to wet forest and subalpine woodland at elevations of 660–3,070 m on Kaua‘i, Moloka‘i, Maui, and Hawai‘i. This species is sister to a species from western North America. The fruit is commonly used in making pies and preserves.

Rubus macraei A. Gray ‘ākala

ENDEMIC | RARE

4.454 

Rubus macraei

Shrubs with shredding bark and prostrate to decumbent stems. The leaves are rather thick and palmately compound, sometimes with scattered small prickles. Fruit dark red, juicy. Uncommon to rare on steep rocky slopes and open rocky areas in wet forest and margins of bogs or alpine shrubland at elevations of 1,610– 2,080 m on East Maui and Hawai‘i.

FLOWERING PLANTS

Rubiaceae

(COFFEE FAMILY)

Herbs, shrubs, trees, or lianas. Key features are simple, opposite leaves, the presence of stipules, the usually 4–5-lobed calyx and corolla, and an inferior ovary. There are 563 genera and about 10,900 species found primarily in the tropics and subtropical regions of the world. Economically the family is important for coffee (Coffea), quinine (Cinchona), and ipecac (Cephaelis). A number of ornamentals, such as gardenia (Gardenia), mirror plant (Coprosma), ixora (Ixora), and needle flower (Posoqueria), are members of this family. In the Hawaiian Islands represented by the endemic genus Bobea and the indigenous genera Coprosma, Gardenia, Gynochthodes, Kadua, Nertera, Psychotria, and Psydrax. At least 12 genera of weedy species have become naturalized, including Paederia, Richardia, Spermacoce, Hedyotis, and Mitracarpus. Coffee is grown commercially in the Hawaiian Islands and has also established itself in the wild.

in cymes borne on peduncles 30–120 mm in length, perfect or functionally unisexual. Occurs in mesic valleys and mesic forest to wet forest at elevations of 260– 1,280 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Restricted to the Ko‘olau Mountains on O‘ahu and Kohala Mountains on Hawai‘i. The wood was formerly used for poi boards and paddles. The gunwales and carved ends of outrigger canoes were also made from this wood because of its bright yellow color and durability.

Bobea elatior Gaudich.

‘ahakea, ‘ahakea lau nui ENDEMIC | SECURE

4.455 

Bobea elatior

Trees or rarely shrubs to 15 m tall. Leaves chartaceous, somewhat coriaceous, elliptic to obovate, 3–15 cm in length. Flowers

Coprosma elliptica W. R. B. Oliv. pilo

ENDEMIC | SECURE

Bobea sandwicensis (A. Gray) Hillebr. ‘ahakea

ENDEMIC | VULNERABLE

4.457 

Bobea—An endemic genus of 4 species of

primarily trees, rarely shrubs. The name honors Jean Bobe-Moreau, a French physician and pharmacist. Leaves entire, opposite, with domatia on lower surface. Flowers functionally unisexual or sometimes perfect, yellow-green and pendulous on long pedicels; fruit drupaceous, purplish to black. The genus is closely related to Timonius of the paleotropics and eastward to Micronesia.

characteristics. In the Hawaiian Islands represented by 13 endemic species, 12 of which have red-orange fruit and 1 with black fruit, indicating at least 2 separate introductions. Some systematists merge Nertera within Coprosma (see below). The fruits of all species are highly prized by native birds.

4.456 

Bobea sandwicensis

Trees to 10 m tall. Leaves chartaceous, somewhat coriaceous, ovate to elliptic, 6.5–11.5 cm in length. Flowers perfect or functionally unisexual, borne in cymes on short peduncles 6–25 mm in length. Occurs in dry to mesic forest and sparsely vegetated lava flows at elevations of 100– 1,220 m on O‘ahu, Moloka‘i, Lāna‘i, and Maui.

Coprosma—A genus of shrubs or small trees, sometimes creeping and often rooting at the nodes. Leaves simple, entire, or dentate on the margins. Flowers unisexual, plants dioecious. Fruit drupaceous and juicy. There are about 100 species primarily in New Zealand, but they are also found in New Guinea, Australia, the Hawaiian Islands, and throughout the Pacific Basin. The name is from the Greek kopros, dung, and osme, fragrant, referring to the fetid smell of the foliage of some species when bruised. The bright orange fruit is highly prized by many of the native birds. Individual species are often separated on the basis of stipule

Coprosma elliptica

Compact or scandent shrubs, nearly prostrate, with many branches, stems 2.0–2.5 m in length, sometimes rooting at the nodes. Leaves opposite, small, 6–10 mm in length, very closely spaced, coriaceous. Dioecious, flowers solitary, fruit yellowish orange. Occurs in open bogs, bog margins, and hummocks from the Alaka‘i Swamp to Mount Wai‘ale‘ale and surrounding wet forest at elevations of 1,190–1,570 m on Kaua‘i.

Coprosma ernodeoides A. Gray

‘aiakanēnē, leponēnē, kūkaenēnē ENDEMIC | SECURE

Trailing shrubs with many prostrate branches and a prostrate growth form. Leaves opposite, very closely spaced, in clusters on short branches, stiff and linear-lanceolate, 6–13 mm in length. Dioecious, flowers solitary. The round, shiny black fruit is distinctive among the Hawaiian species of Coprosma. Occurs on lava and cinder fields in subalpine woodland at elevations of 1,220–2,590 m on West Maui (Mount ‘Eke, East Bog), East Maui, and Hawai‘i (see figure 2.8). This species represents a separate colonizing lineage from the other Hawaiian species. The fruit is eaten by the native Hawaiian goose.

145

146

FLOWERING PLANTS

Coprosma foliosa A. Gray pilo

ENDEMIC | SECURE

Shrubs or small trees 2–6 m tall, stems pubescent. Leaves opposite, chartaceous, elliptic to spatulate, 4–12 cm in length. Dioecious, flowers typically 3 in a cluster, fruit bright reddish orange. Occurs in mesic to wet forest at elevations of 600– 1.330 m on Kaua‘i. Closely related to C. ochracea.

Coprosma longifolia A. Gray pilo

ENDEMIC | SECURE

Diffusely branched to scandent shrubs, stems 2–4 m or more in length. Leaves opposite, elliptic to ovate, 1.8–6.0 cm in length. Dioecious, flowers in clusters of 1–4, fruit bright orange to yellow orange. Occurs in mesic forest at elevations of 270–1,220 m on Hawai‘i. Closely related to the widespread and polymorphic C. foliosa.

Coprosma montana Hillebr. pilo

ENDEMIC | SECURE

4.458 

Coprosma foliosa

Scandent shrubs to small trees, 1–8 m in height, stems with many branches. Leaves opposite, typically closely spaced but widely so in scandent plants, charta­ ceous, lanceolate to elliptic or obovate, 2–11 cm in length. Dioecious, flowers 1–3 per cluster, fruit bright to reddish orange. Occurs in mesic to wet forests or occasionally ridges at elevations of 300– 1,830 m on Kaua‘i (rare), O‘ahu, Moloka‘i, Lāna‘i, and Maui.

Coprosma kauensis (A. Gray) A. Heller pilo

ENDEMIC | SECURE

4.460 

Coprosma longifolia

Shrubs or small trees 2–5 m tall, young stems glabrous. Leaves in whorls of 3, thick, chartaceous, lanceolate to oblong or elliptic-lanceolate, 6.5–11.5 cm in length. Dioecious, flowers typically in clusters of 6–12, fruit reddish orange. Occurs in mesic to occasionally wet forest on both mountain ranges on O‘ahu at elevations of 360–1,200 m. Male flowers are pictured.

Coprosma menziesii A. Gray pilo

ENDEMIC | SECURE

4.459 

Coprosma kauensis

4.461 

Coprosma menziesii

4.462 

Coprosma montana

Many-branched small trees or shrubs with stiff, ascending branches. Leaves opposite, thin to coriaceous in texture, often clustered near branch tips, oblanceolate to broadly elliptic-ovate, 1–5 cm in length. Dioecious with notable sexual dimorphism in shrub architecture, flowers in clusters of 1–2, fruit orange to reddish orange. Occurs in subalpine shrub­ land, where it is often a dominant, or in subalpine woodland to mesic forest at elevations of 1,830–3,050 m on Moloka‘i (Pu‘u Ali‘i Natural Area Reserve), East Maui, and Hawai‘i.

FLOWERING PLANTS

Coprosma ochracea W. B. R. Oliv. pilo

ENDEMIC | SECURE

fruit bright orange to reddish orange. Occurs in wet forest or less commonly in mesic forest at elevations of 360–1,900 m on Moloka‘i, Maui, and Hawai‘i.

Coprosma ternata W. R. B. Oliv. pilo

ENDEMIC | SECURE

4.463 

Coprosma pubens A. Gray pilo

ENDEMIC | SECURE

Gardenia brighamii H. Mann nānū, nā‘ū

ENDEMIC | ENDANGERED

4.465 

Coprosma ternata

Shrubs 2–5 m tall with ascending to spreading stems, stems glabrous. Leaves in whorls of 3, elliptic to oblanceolate, somewhat thick and chartaceous, 5.0– 10.5 cm in length. Dioecious, flowers in clusters of 10–18 or more, fruit reddish orange. Occurs in mesic to wet forest at elevations of 760–1,400 m on eastern Moloka‘i and West Maui (Hana‘ula). Known to hybridize with C. ochracea.

Coprosma waimeae Wawra ‘ōlena, pilo

ENDEMIC | SECURE

4.464 

Coprosma pubens

Shrubs or small trees 3–10 m tall, stems many branched, typically glabrous. Leaves opposite, thin, coriaceous, oblong elliptic to broadly obovate, 5.0–13.5 cm in length. Dioecious, flowers 3–4 per cluster,

Gardenia—A paleotropical and warm

temperate Asian genus of trees and shrubs represented by about 200 species. Young branches often covered with a sticky exudate from the stipules. Flowers solitary and terminal; fruit globose and large. The name honors Alexander Garden, an eighteenth-century correspondent of Linnaeus. In the Hawaiian Islands, there are 3 endemic species, 2 of which are quite rare. These species represent 2 separate colonizing lineages. A few species are cultivated for their fragrant white flowers.

Coprosma ochracea

Erect shrubs or small trees 2–6 m tall or in bogs with a prostrate or weakly spreading growth form, many branched, lateral branches in a single flat plane in shrubs and bog forms, stems pubescent. Leaves opposite, coriaceous, variable in shape from elliptical to broadly ovate, 1.5–10.0 cm in length. Dioecious, flowers 3–6 per cluster, fruit bright orange. Occurs in wet forest and bogs and occasionally mesic forest at elevations of 790–2,290 m on O‘ahu (Mount Ka‘ala) and on eastern Moloka‘i, Lāna‘i, Maui, and Hawai‘i. A polymorphic species often hybridizing with other species of Coprosma where they are sympatric.

Shrubs 1.5–4.0 m tall. Leaves opposite, thin, coriaceous, elliptic to obovate, 6.0– 11.5 cm in length. Dioecious, flowers 1–4 per cluster, fruit bright orange. Occurs in mesic forest at elevations of 580–1,070 m on Kaua‘i.

4.466 

Coprosma waimeae

4.467 

Gardenia brighamii

Small trees up to 5 m tall, branches 4 angled in cross section. Leaves elliptic, 2.2–10.5 cm in length, few in number, and clustered toward the tips of the branches. Flowers white and fragrant, solitary. A dry forest species virtually extinct in the wild. A few extant plants had survived in dry forest remnants at elevations of 350–520 m on O‘ahu (Wai‘anae Mountains: Pu‘uku‘ua, Nanakuli), Moloka‘i (Mauna Loa, Mahana Flats, extinct), Lāna‘i (Kānepu‘u), West Maui (Olowalu, extinct), and Hawai‘i (Pu‘uwa‘awa‘a). Maintained in cultivation in botanical gardens in the Hawaiian Islands. Extremely rare, with estimates that the total extant population is 15–19 trees, with 2 in the wild on O‘ahu and 1 on Hawai‘i. Threats have come from dry forest habitat loss, alien plant species, and goats.

147

148

FLOWERING PLANTS

Gynochthodes—A genus of 93 species of

lianas or rarely shrubs or trees. Leaves petiolate and leathery in texture. Flowers polygamous or dioecious. Distributed in tropical Asia, Australasia, and Madagascar, with a single endemic species in the Hawaiian Islands. The name comes from the Greek gyn, female, and ochthodes, warty, referring to the warty surface of the stigma. This species was placed in the genus Morinda in the Manual.

and 1 widely distributed Pacific species. Several species with fleshy fruit were once segregated as the endemic genus Gouldia. The Hawaiian taxa represent a radiation from a single founding lineage and include both rare and extinct species.

Kadua acuminata Cham. & Schltdl. au, pilo

ENDEMIC | SECURE

Gynochthodes trimera (Hillebr.) Razafim. & B. Bremer noni kuahiwi

ENDEMIC | SECURE

Shrubs, lianas, or small trees to 5 m tall. Leaves highly variable, with textures from membranaceous to chartaceous to coriaceous, shapes from ovate to elliptic or suborbicular, 2–18 cm in length, flat to weakly concave. Inflorescences also highly variable in form, typically terminal or on short leafy branches. Occurs in mesic to wet forest, bogs, and mesic shrubland at elevations of 260–2,040 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Next to Metrosideros polymorpha, this is arguably the most polymorphic species among the flowering plants in the Hawaiian Islands, with dozens of infraspecific taxa named in the past. Formerly known as Gouldia terminalis and listed in the Manual as Hedyotis terminalis.

Kadua axillaris (Wawra) W. L. Wagner & Lorence manono 4.469 

4.468 

Gynochthodes trimera

Trees 4–16 m tall, branches terete or sometimes 4 angled, with many warts and lenticels. Leaves chartaceous, upper surface glossy, oblanceolate to elliptic, 12–26 cm in length. Flowers are borne in an irregular rounded head that later forms a woody conglomerate fruit. This is 1 of only 2 arborescent species in the genus. Occurs in mesic forest at elevations of 300–880 m on Kaua‘i, O‘ahu, Lāna‘i, and Maui. Listed in the Manual as Morinda trimera.

Kadua—A genus of 29 species of herbs,

shrubs, subshrubs, small trees, or occasionally herbaceous perennials or lianas; endemic to Polynesia, with a major node of speciation in the Hawaiian Islands. Flowers perfect or functionally pistillate and perhaps sometimes functionally staminate, fruit a capsule. The name honors M. Kadu, a member of the exploring expedition of Otto von Kotzebue in the early nineteenth century. These species were formerly included in Hedyotis in the Manual. In addition to 22 endemic species in Hawaii, Kadua includes 4 species from the Marquesas, 2 species from the Society Islands, 1 species from Rapa Nui,

Kadua acuminata

ENDEMIC | SECURE

Shrubs or subshrubs, stems 2–4 m in length with lateral branches 10–100 cm in length, stems with a fetid smell when bruised. The widely spaced leaves are variable in shape from lanceolate to ovate, chartaceous to somewhat coriaceous, 7–18 cm in length. Similar to K. fluviatilis but flowers yellowish green, green, or greenish white and generally smaller. Occurs in mesic to occasionally wet forest at elevations of 90–800 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in the Manual as Hedyotis acuminata.

Kadua affinis DC. manono

ENDEMIC | SECURE

4.471 

4.470 

Kadua affinis

Kadua axillaris

Shrub to small tree 2–6 m tall, occasionally trailing as a liana. Leaves evenly spaced, membranous to chartaceous, oblanceolate to obovate, 6–16 cm in length. Paniculate inflorescences axillary or on very short leafless branches, flowers perfect, corolla greenish white. Occurs in mesic to wet forests at elevations of 40–1,830 m on Moloka‘i, Maui, and Hawai‘i. Listed in the Manual as Hedyotis hillebrandii.

FLOWERING PLANTS

Kadua centranthoides Hook. & Arn. ENDEMIC | SECURE

occurred at Kealakekua, Hawai‘i. Listed in the Manual as Hedyotis cookiana. Recent surveys have found only 2 extant populations with a total of 100–122 individuals, all on Kaua‘i. Threats come from feral pigs, goats, alien plant species, and landslides.

Kadua cordata Cham. & Schltdl. kopa

ENDEMIC | SECURE/ENDANGERED

Loose, branched subshrubs that often form clumps or mats, stems weakly ascending 10–20 cm in length, weakly terete to 4 angled. Leaves thin, linear-elliptic or lanceolate, 4–18 cm in length, margins flat to slightly revolute. Flowers perfect and pistillate, corolla white to greenish, occasionally pale purple. Uncommon in wet sites such as cliff faces, streambeds, and waterfalls at elevations of 120–1,100 m on Kaua‘i, O‘ahu, Moloka‘i, and East Maui. Listed in the Manual as Hedyotis elatior.

Kadua fluviatilis C. N. Forbes kamapua‘a, pilo

ENDEMIC | RARE

4.472 

Kadua centranthoides

Erect to sprawling subshrubs, stems 30–200 cm in length, terete, leaves widely spaced, coriaceous, variable in shape, 6–16.5 cm in length. Lateral veins typically impressed on the upper surface. Inflorescence elongate, flowers perfect or functionally pistillate, corolla purple to violet. Occurs in wet forest and along the margins of bogs and sometimes in mesic forest at elevations of 380–1,920 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in the Manual as Hedyotis centranthoides.

Kadua cookiana Cham. & Schltdl. ‘āwiwi

ENDEMIC | ENDANGERED

4.474 

Kadua cordata subsp. cordata

Weakly scandent subshrubs, stems with few branches. Leaves glossy, thin and membranous, variable in shape from ovate to elliptic and occasionally lanceolate, 3–16 cm in length, veins slightly to strongly impressed on upper surface. Flowers perfect and pistillate, corolla pale green, greenish yellow, or cream. Occurs in mesic to wet forest at elevations of 400– 1,820 m primarily on Kaua‘i and O‘ahu but also on Moloka‘i, Lāna‘i, and Maui. A polymorphic species with 3 subspecies recognized. The subsp. cordata shown here is widely distributed on Kaua‘i, O‘ahu, Moloka‘i, and Maui. The subsp. waimeae is restricted to Kaua‘i, while the endangered subsp. remyi occurs on Lāna‘i. Listed in the Manual as Hedyotis schlechtendahliana.

Kadua elatior (H. Mann) W. L. Wagner & Lorence ‘āwiwi ENDEMIC | SECURE

4.476 

Kadua fluviatilis

Scandent shrubs, stems 30–250 cm in length with short lateral branches that have a fetid smell when bruised. Leaves widely spaced and variable in shape from ovate to elliptic-lanceolate, 8–17 cm in length. Flowers perfect and pistillate, corolla white. Similar to K. acuminata but differs in having waxy white flowers up to 20 mm in length. A rather rare species in mesic to wet forest at elevations of 60–1,220 m on Kaua‘i and in the Ko‘olau Mountains (Pūpūkea to Mānoa), O‘ahu. Listed in the Manual as Hedyotis fluviatilis.

Kadua foggiana (Fosberg) W. L. Wagner & Lorence ENDEMIC | SECURE

4.473 

Kadua cookiana

Small multibranched shrub with stems up to 20 cm in length. Leaves chartaceous, thin, linear, 4–8 cm in length. Upper parts of plant and inflorescence notably glaucous. Flowers perfect or pistillate, corolla white. Very rare, with only a few extant populations, although thought to have once been more common. Known from lowland habitats in the Waiahuakua Valley and Honakoa, Kaua‘i, and formerly

4.475 

Kadua elatior

4.477 

Kadua foggiana

149

150

FLOWERING PLANTS

Sprawling to scandent subshrub with diffuse branches 1–4 m in length. Leaves widely spaced, coriaceous, elliptic to ovate in shape, 6.5–16.0 cm in length. Inflorescence broad and diffuse. Flowers perfect or functionally pistillate, corolla green to greenish yellow. Occurs in mesic to wet forests at elevations of 750–1,220 m on Kaua‘i (Mount Kāhili, Hanalei Valley, Kōke‘e). Listed in the Manual as Hedyotis foggiana.

Diffuse, scandent shrubs with a weeping habit, stems 100–400 cm in length. Leaves widely spaced, chartaceous, ovate to broadly elliptic, 6–10 cm in length. Flower perfect and pistillate, corolla green. Occurs in mesic forest at elevations of 610–1,100 m in the Kōke‘e area on Kaua‘i. Listed in the Manual as Hedyotis knudsenii.

Kadua formosa Hillebr.

ENDEMIC | VULNERABLE

Kadua tryblium (D. R. Herbst & W. L. Wagner) W. L. Wagner & Lorence ENDEMIC | RARE

Kadua littoralis Hillebr.

ENDEMIC | RARE

4.481  4.478 

Kadua formosa

Subshrubs up to 1 m tall with the main stem prostrate, stems somewhat 4 angled. Leaves coriaceous, lanceolate to elliptic, 5.5–14.0 cm in length. Corolla white. A rare plant that occurs on wet cliffs on West Maui. Listed in the Manual as Hedyotis formosa.

Kadua knudsenii Hillebr. ENDEMIC | SECURE

4.480 

4.479 

Kadua knudsenii

Kadua littoralis

Succulent perennial herbs, slightly woody at their base, basal leafy stems 25–40 cm in length. Leaves clustered at the base of the stems, fleshy, upper surface glossy, ovate to broadly elliptic or sometimes lanceolate, 3.5–8.5 cm in length. Flowers perfect, corolla white. Occurs on wet sea cliffs and coastal talus slopes, usually within the zone of salt spray, on Kaua‘i (Princeville), O‘ahu (extinct), windward Moloka‘i, East Maui (Ke‘anae-Wailua, Hanawī, ‘Ālau Island), and Hawai‘i (Honopue Valley). Listed in the Manual as Hedyotis littoralis.

Kadua tryblium

Scandent shrubs, stems trailing up to 5 m in length with few branches, often rooting when in contact with the soil. Leaves thick, green above and pale green beneath with purplish splotches, elliptic in shape and slightly asymmetrical, 13–22 cm in length. The unusual stipules adhere to the petioles, forming a small cup otherwise unknown in any of the other Hawaiian Kadua. Flowers perfect or functionally pistillate, corolla yellowish green. A relatively recently described species from wet forest on Kaua‘i (Mount Kāhili, Wahiawa Stream, rim of Kalalau Valley). Listed in the Manual as Hedyotis tryblium.

Morinda—Morinda trimera, listed in the

Manual, has now been placed into the genus Gynochthodes.

Nertera—A genus of 6 species of trailing perennial herbs with orange fruit in Malesia, China to Taiwan, New Zealand, Polynesia, temperate South America, and Tristan de Cunha (Atlantic Ocean). Some systematists merge Nertera into Coprosma. The name is from the Greek nerteros, lowly, because of the creeping habit of the plants. In the Hawaiian Islands represented by 1 indigenous species.

FLOWERING PLANTS

Nertera granadensis (Mutis ex L. f.) Druce mākole

INDIGENOUS | SECURE

4.482 

Nertera granadensis

Perennial herbs with prostrate branches that root at the nodes, often forming mats. The leaves are slightly fleshy. Distributed widely in South America, the Juan Fernandez Islands, and Indonesia. Occurs in the Hawaiian Islands on wet soil, sometimes epiphytic on logs in wet forest at elevations of 490–2,070 m on Kaua‘i, O‘ahu (Wai‘anae Mountains), Moloka‘i, Maui, and Hawai‘i. Tissues hyperaccumulate manganese.

Trees to 12 m tall. Leaves chartaceous to coriaceous, elliptic to orbicular, glabrous to pubescent, domatia in the vein axils. Flowers functionally unisexual. Three varieties are recognized that differ primarily in their leaf pubescence. The var. hillebrandii, pictured, occurs in wet forest and occasionally dry to mesic forest at elevations of 150–1,590 m on Moloka‘i, Maui, and most commonly Hawai‘i. The var. hawaiiensis occurs on Moloka‘i, Maui, and Hawai‘i, while the var. scoriacea is restricted to Maui and Hawai‘i.

Psychotria hawaiiensis (A. Gray) Fosberg kōpiko ‘ula, ‘ōpiko

Psychotria kaduana (Cham. & Schltdl.) Fosberg kōpiko kea

ENDEMIC | SECURE

Psychotria hexandra H. Mann ‘ōpiko

ENDEMIC | SECURE/RARE

4.486 

Psychotria—A large genus of 1,850 or more

species found in both the neotropics and paleotropics. Flowers perfect but usually functionally unisexual, borne in panicles; fruit drupaceous. Leaves often with domatia. Straussia, once considered an endemic Hawaiian genus, is placed here. The name is from the Greek psychros, cold, and trophos, feeder. Linnaeus derived the current name from psychotrophon, a name applied to a Jamaican plant. In the Hawaiian Islands represented by 11 endemic species, which are the result of 2 separate colonizing lineages.

Trees to 8 m tall. Leaves membranous, green above and rose colored or pink on the lower surface, domatia small or absent. Flowers functionally unisexual. Occurs in mesic forest in a very localized area at elevations of 600–610 m in western Kaua‘i (Miloli‘i-Kopiwai). One of the most ornamental species and worthy of cultivation.

4.484 

Psychotria hexandra var. hexandra

Trees or shrubs up to 6 m tall. Leaves membranous to chartaceous or coriaceous, obovate to oblanceolate, domatia small and inconspicuous. Flowers unisexual. Occurs in mesic to wet forest at elevations of 360–1,250 m on Kaua‘i and O‘ahu. There are 2 named varieties, with var. hexandra on Kaua‘i and the rare var. oahuensis in the Ko‘olau Mountains.

Psychotria kaduana

Trees and shrubs 2–4 m tall or more with obovate to elliptic leaves. The flowers are often in a pendulous inflorescence. Flowers functionally unisexual. Occurs in mesic valleys, mesic forest, and wet forest at elevations of 180–1,220 m Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui. This is the most widely distributed species in the Islands.

Psychotria mariniana (Cham. & Schltdl.) Fosberg ‘ōpiko ENDEMIC | SECURE

Psychotria hobdyi Sohmer kōpiko

ENDEMIC | ENDANGERED

ENDEMIC | SECURE

4.487 

4.483 

Psychotria hawaiiensis var. hillebrandii

4.485 

Psychotria hobdyi

Psychotria mariniana

Trees up to 20 m tall. Leaves coriaceous, narrowly oblanceolate to obovate,

151

152

FLOWERING PLANTS

typically glabrous, domatia conspicuous. Flowers functionally unisexual. A variable species in morphology and habitat of occurrence. Occurs in dry to wet forests at elevations of 180–1,220 m on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui.

Psydrax odorata (G. Forst.) A. C. Sm. & S. Darwin alahe‘e, ‘ōhe‘e, walahe‘e INDIGENOUS | SECURE

Melicope—A genus of trees and shrubs

Psychotria mauiensis Fosberg ‘ōpiko

ENDEMIC | SECURE

4.489 

4.488 

Psychotria mauiensis

Variable in architecture from shrubs 4 m in height to trees 12 m tall. Leaves membranaceous to coriaceous, broadly ovate to elliptic or suborbicular, domatia absent or inconspicuous. Flowers functionally unisexual. A highly variable species in leaf shape and size, pubescence, and size of the floral organs. Occurs in mesic to wet forests at elevations of 215–1,470 m on Kaua‘i, Moloka‘i, Lāna‘i, Maui, and Hawai‘i.

Psydrax—A genus of shrubs or small trees

closely related to Canthium, with about 75 paleotropical species found in Africa, India, Ceylon, China, Australia, and the Pacific including the Hawaiian Islands. Leaves coriaceous and often shiny on their upper surface; flowers perfect or unisexual. The name is from the Greek psydrax, blister, referring to the wrinkled appearance of the fruit of some species. In the Hawaiian Islands represented by 1 widespread indigenous species.

Zanthoxylum. Species of Melicope and Platydesma in the Hawaiian Islands represent a single founding lineage.

Psydrax odorata

Shrubs or small trees with glossy, somewhat coriaceous leaves that are variable in shape from broadly elliptic and ovate to lanceolate. The white flowers, borne in dense inflorescences, are very fragrant. Native to Micronesia and the South Pacific from Vanuatu to New Caledonia and east to the Tuamotus and the Hawaiian Islands. Occurs in dry shrubland, dry to mesic forest, and sometimes in wet forest at elevations of 10–860 m or higher on all of the main islands except Ni‘ihau and Kaho‘olawe. The dense wood was used for tools and adz blades for cutting softer wood. A black dye was reputed to have been produced from the leaves. Listed in the Manual as Canthium odoratum.

found in the paleotropics from Polynesia to Australasia, China, India, and Madagascar. Foliage and flower parts usually glandular punctuate, typically with waxy scales on young foliage. Flowers in most species functionally unisexual, and the plants are polygamous. The fruit are 4-lobed capsules, often with 1 of the follicles aborted, and may be cuboid or cruciform in shape. There are 233 species, of which 48 are endemic to the Hawaiian Islands. Studies have indicated that Pelea, to which the Hawaiian and Marquesan species were previously assigned, should be merged into Melicope. Many of the Hawaiian plants are highly variable and in need of study. Recent molecular data indicate that Platydesma is nested within Melicope and has evolved from the same colonizing lineage, and thus it should not represent a distinct endemic genus.

Melicope anisata (H. Mann) T. G. Hartley & B. C. Stone alani, alani kuahiwi, mokehana ENDEMIC | SECURE

Rutaceae

(CITRUS FAMILY)

A family of trees, shrubs, woody lianas, and herbs. Key features are the presence of translucent pellucid oil glands on the foliage, 2 whorls of stamens with the outer one opposite the petals, and a lobed ovary raised up on a disc. The oil glands are the source of the characteristic citrus odor. There are 158 genera and about 1,900 species found primarily in tropical or warm temperate regions of the world but also present in arid regions. Economically the family is important for the number of citrus fruits (lemons, oranges, grapefruit, limes, and others). Among the cultivated ornamentals are the cape chestnut (Calodendrum), rue (Ruta), orange jessamine (Murraya), dittany (Dictamnus), and prickly ash (Zanthoxylum). In the Hawaiian Islands represented by 1 endemic genus, Platydesma, and 2 indigenous genera, Melicope and

4.490 

Melicope anisata

Shrubs or sometimes shrublike trees 2–8 m tall, with lax stems with few branches. Leaves opposite, typically 10–20 cm in length, thin, coriaceous, elliptic to oblong-elliptic, with a strong smell of anise when crushed. Separate male and female flowers. Fruit cuboid in shape and also strongly anise scented. Occurs in diverse mesic to wet forest at elevations

FLOWERING PLANTS

elliptic, chartaceous to coriaceous. Fruit cruciform. Occurs in bogs and wet forest at elevations of 520–1,490 m on Kaua‘i. Listed in the Manual as Pelea feddei.

of 370–1,220 m on Kaua‘i. The twigs and fruit were formerly used by the Hawaiians to scent their kapa cloth. The fruit is used in making lei. Listed in the Manual as Pelea anisata.

Melicope hawaiensis (Wawra) T. G. Hartley & B. C. Stone

Melicope barbigera A. Gray

alani, alani kuahiwi, uahiapele

alani, alani kuahiwi, manena, mokihana kūkae moa

ENDEMIC | SECURE

ENDEMIC | VULNERABLE

4.493 

4.491 

Melicope barbigera

Small trees 3–12 m tall. Leaves dull green, 8–20 cm or more in length, weakly scented, with soft hairs in a line along each side of the midrib on their lower surface Common in mesic forest at elevations of 850–1,220 m in the Kōke‘e area on Kaua‘i. Listed in the Manual as Pelea barbigera.

Melicope clusiifolia (A. Gray) T. G. Hartley & B. C. Stone

kolokolo mokihana, kūkaemoa, alani, alani kuahiwi ENDEMIC | SECURE

Melicope clusiifolia—O‘ahu

Trees or shrubs 2–10 m tall. Leaves typically in whorls of four, 5–30 cm in length, variable in shape, more or less coriaceous. The flowers perfect or functionally unisexual and then the plants dioecious, maturing on naked stems below the leaves. This is the most widespread and variable species of Melicope in the Hawaiian Islands. Common in wet forest to mesic forest on all of the main islands except Ni‘ihau and Kaho‘olawe. Shown here are plants from Kaua‘i and O‘ahu, each with distinctive morphological traits. The shiny black seeds are eaten by birds. Listed in the Manual as Pelea clusiifolia.

Melicope feddei (H. Lev.) T. G. Hartley & B. C. Stone alani

4.495 

Melicope hawaiensis

Shrubs or trees 3–10 m tall. Leaves opposite, 5.5–20.0 cm in length, coriaceous, elliptic, lower surface sparsely to densely hairy beneath. Fruit somewhat cruciform, covered with golden brown or reddish brown hairs. Occurs in dry forest and occasionally mesic forest at elevations of 610–1,220 m on Moloka‘i, Lāna‘i, Maui, and Hawai‘i. A widespread and variable species. Listed in the Manual as Pelea hawaiensis.

Melicope kaalaensis (H. St. John) T. G. Hartley & B. C. Stone alani ENDEMIC | SECURE

ENDEMIC | SECURE

4.496 

4.494 

4.492 

Melicope clusiifolia—Kaua‘i

Melicope feddei

Sprawling, prostrate, or erect shrubs 1–2 m tall, with reddish brown branches. Leaves opposite, 2–8 cm in length,

Melicope kaalaensis

Small to medium-sized trees 3–7 m tall. New growth with dense yellowish pubescence. Leaves with a strong anise odor, opposite, coriaceous, ovate to elliptic, 4–23 cm in length, upper surface glossy dark green, lower surface paler with

153

154

FLOWERING PLANTS

reddish brown veins. Separate male and female flowers. Fruit cuboid, green. Occurs in mesic forest at elevations of 520–860 m in the Wai‘anae Mountains (Pu‘ukaua to Piko Trail), O‘ahu. Listed in the Manual as Pelea kaalaensis.

Melicope peduncularis (H. Lév.) T. G. Hartley & B. C. Stone alani ENDEMIC | SECURE

Melicope makahae (B. C. Stone) T. G. Hartley & B. C. Stone alani

ENDEMIC | ENDANGERED

4.499 

Melicope oahuensis—fruit

Shrubs to shrubby trees 2–7 m tall. Leaves opposite, typically 6–12 cm in length, thin, coriaceous, elliptic, with a strong anise odor when crushed. Flowers functionally unisexual or perfect, with both staminate and perfect flowers on the same plant. The fruit is cube shaped with a central depression. Occurs in mesic to occasionally wet forest at elevations of 430–880 m or higher on O‘ahu. Listed in the Manual as Pelea oahuensis.

Melicope ovata (H. St. John & E. P. Hume) T. G. Hartley & B. C. Stone alani ENDEMIC | SECURE

4.497 

Melicope makahae

Shrubs or shubby trees to 3 m tall. Leaves opposite, 9.5–12.0 cm in length, coriaceous, ovate to elliptic. Separate male and female flowers. Fruit dark green. Occurs in mesic forest at elevations of 670–1,220 m in the Wai‘anae Mountains (Pālāwai Gulch to Mākaha Valley, ridges), O‘ahu. Listed in the Manual as Pelea makahae.

4.501 

Melicope peduncularis

Erect to sprawling shrubs or shrubby trees, 1–4 m tall. Leaves opposite or in whorls of 3–4 per node, 3–23 cm in length, elliptic to suborbicular, dark green on the upper surface, coriaceous, with a strong anise odor when crushed. Separate male and female flowers. Fruit cuboid. Occurs in mesic forest at elevations of 300–1,160 m on Kaua‘i (Kōke‘e), O‘ahu, eastern Moloka‘i (Ka‘aloa Peak), and Maui. Listed in the Manual as Pelea peduncularis.

Melicope puberula (H. St. John) T. G. Hartley & B. C. Stone alani

ENDEMIC | RARE

Melicope oahuensis (H. Lev.) T. G. Hartley & B. C. Stone alani, alani kuahiwi ENDEMIC | SECURE

4.500 

4.498 

Melicope oahuensis—flower

Melicope ovata

Shrubs to small trees 3–8 m tall, branches sprawling. Leaves opposite, 7–20 cm in length, thin, coriaceous to chartaceous, Separate male and female flowers. Fruit cruciform. Occurs primarily in mesic forest at elevations of 55–1,070 m on northwest Kaua‘i and the Wai‘anae Mountains (Napepeiaulelo Gulch, North Pālāwai Gulch), O‘ahu. Listed in the Manual as Pelea ovata.

4.502 

Melicope puberula

FLOWERING PLANTS

Shrubs or small trees, 3–10 m tall, young stems with dense yellow-brown pubescence. Leaves opposite, coriaceous, narrowly elliptic to elliptic ovate, 6–17 cm in length, lower surface sparsely to moderately pubescent. Separate male and female flowers. Fruit cuboid, dark red when mature. Rare in mesic to wet forest at elevations of 1,070–1,220 m from Kalalau Lookout to Pu‘uokila Lookout, Kaua‘i. Recent surveys have reported 3 extant populations totaling about 900 individuals. Listed in the Manual as Pelea puberula.

Melicope sessilis (H. Lév.) T. G. Hartley & B. C. Stone alani

1,220–1,590 m on Kaua‘i (Wahiawa Bog, Alaka‘i Swamp to Mount Wai‘ale‘ale). Listed in the Manual as Pelea waialealae.

ENDEMIC | SECURE

Platydesma—An endemic Hawaiian genus of 4 species of shrubs or small trees. Often sparingly branched, giving them a palmlike form of growth. Traditionally separated from Melicope by the presence of perfect flowers and stamens joined in a filament tube rather than distinct in 2 whorls. The name is from the Greek platys, flat, and desmos, chain, referring to the form of the filaments of the anthers. The plants have a smell reminiscent of stomach gas when bruised or broken. Recent molecular studies indicate that Platydesma is nested within Melicope and thus does not represent a distinct endemic genus.

Melicope rotundifolia (A. Gray) T. G. Hartley & B. C. Stone alani

ENDEMIC | SECURE

4.504 

Melicope sessilis

Shrubs to 3 m tall. Leaves crowded, opposite, 4–10 cm in length, elliptic, with short petioles. Separate male and female flowers borne on slender pedicels. Fruit cuboid. Occurs in wet forest at elevations of 1,000–1,400 m on Moloka‘i (Waikolu to Kaunohua), West Maui (Honokōwai to Pu‘ukukui), and East Maui (Olinda to ‘Ōpana Gulch). Listed in the Manual as Pelea parvifolia.

Platydesma cornuta Hillebr. ENDEMIC | RARE

Melicope waialealae (Wawra) T. G. Hartley & B. C. Stone alani, alani kuahiwi, ‘ānonia, alani wai ENDEMIC | SECURE

4.506  4.503 

Melicope rotundifolia

Scandent shrubs with stems 1–3 m in length. Leaves opposite or rarely in whorls of 3, 6–13 cm in length, coriaceous, ovate to suborbicular, margins strongly inrolled. Separate male and female flowers. Fruit cruciform. Occurs in mesic forest to wet forest at elevations of 360–760 m in the Ko‘olau Mountains (‘Ōpae‘ula to Niu Valley), O‘ahu. Listed in the Manual as Pelea rotundifolia.

4.505 

Melicope waialealae—habit

Many-branched shrubs or small trees 0.5–3.0 m tall, stems erect to nearly prostrate in bogs. Leaves in whorls of 4 per node, 5–9 cm in length, coriaceous, narrowly elliptic. Separate male and female flowers. Fruit cuboid. Occurs in open bogs and bog hummocks at elevations of

Platydesma cornuta var. decurrens

Erect, palmlike shrubs 1–2 m tall but occasionally as tall as 7 m. Leaves opposite, 12–40 cm in length, narrowly obovate and clustered toward the end of the branches. Fruit cruciform with 4 hornlike projections. Occurs in mesic forest at elevations of 340–920 m on O‘ahu. Two varieties are recognized, with var. decurrens from mesic forest in the Wai‘anae Mountains pictured. The var. cornuta, with shorter leaf bases, occurs in the Ko‘olau Mountains.

155

156

FLOWERING PLANTS

Platydesma rostrata Hillebr.

Platydesma spathulata (A. Gray) B. C. Stone

ENDEMIC | RARE

ENDEMIC | SECURE

pilo kea lau li‘i

pilo kea

4.509 

4.507 

Platydesma rostrata—habit

Platydesma spathulata

Shrubs or shrubby trees 2–4 m or even 12 m tall, with spreading branches. Leaves opposite, 10–50 cm in length, coriaceous, spathulate to oblanceolate. Fruit cruciform with 4 lobes without projections. Occurs in mesic to wet forest at elevations of 450–1,420 m on Kaua‘i, O‘ahu, Maui, and Hawai‘i. This is the most common member of the genus.

Trees 4–15 m tall with trunks up to 30 cm diameter, wood yellowish. Leaves compound with 3–7 leaflets, coriaceous, the lowest leaflet pair typically reduced in size. The rare var. dipetalum has glabrous leaves and occurs in dry to mesic forest or on the margins of wet forest at elevations of 260–1,280 m on Kaua‘i, O‘ahu, Moloka‘i, and Hawai‘i. The endangered var. tomentosum, with heavily pubescent leaves, is confined to lava fields in North Kona, Hawai‘i. This species represents a separate colonizing lineage from the other 3 species of Zanthoxylum. The trunk was used by Native Hawaiians for kapa logs and anvils. The wood also has resonant qualities and was used as a beater for long-distance drumming.

Zanthoxylum hawaiiense Hillebr. a‘e, hea‘e, mānele

ENDEMIC | ENDANGERED

Zanthoxylum—A large genus of evergreen and deciduous trees and shrubs and sometimes climbers; stems usually spiny but sometimes unarmed. Pantropical in distribution with about 200 species, including some found in temperate regions. The name is from the Greek xanthos, yellow, and xylon, wood, referring to the color of the wood of many of these plants. In the Hawaiian Islands, all 4 of the endemic species are unusual in that they lack prickles or thorns. These species represent 2 separate colonizing lineages.

Zanthoxylum dipetalum H. Mann kāwa‘u, a‘e, hea‘e, mānele ENDEMIC | RARE

4.508 

Platydesma rostrata

Erect, palmlike shrubs 1–3 m tall, with a few ascending stems. Leaves opposite, 20–42 cm in length, narrow oblong to elliptic, clustered toward the ends of the branches. Fruit cruciform, with 4 slender pointed beaks. Rare in diverse mesic forest and mesic valleys at elevations of 760– 1,220 m on Kaua‘i.

4.510 

Zanthoxylum dipetalum var. dipetalum

4.511 

Zanthoxylum hawaiiense

Small to medium-sized trees 3–8 m tall, with trunks up to 25 cm diameter. Leaves compound with 3 coriaceous leaflets, broadly deltate to lanceolate, the lateral ones asymmetrical. Occurs in dry forest at elevations of 550–1,740 m on Kaua‘i, Moloka‘i, Lāna‘i (extinct), Maui, and Hawai‘i. This is the least common species of the genus in the Hawaiian Islands. Past surveys have reported 11 extant populations with a total of about 250 individuals on the four islands where the species survives. Threats come from feral and domestic grazing animals and alien plant species.

FLOWERING PLANTS

Zanthoxylum kauaense A. Gray

Xylosma hawaiiense Seem.

ENDEMIC | SECURE

ENDEMIC | SECURE

a‘e, hea‘e, mānele

4.512 

maua

4.513 

Xylosma hawaiiense—habit, female plant

Salicaceae

(WILLOW FAMILY)

A family traditionally limited to a small number of genera and species of temperate forest and riparian trees, with Salix (willows) and Populus (cottonwoods and aspen) as the primary groups. New molecular data have greatly expanded the concept of the family, and the Salicaceae now includes a number of tropical and Southern Hemisphere genera that were previously included in the Flacourtiaceae. In this expanded concept, there are 54 genera and about 1,200 species. These include Xylosma, which is present in the Hawaiian Islands.

Xylosma—A genus of 85 species of mostly dioecious trees and shrubs, some spiny. They are almost pantropical in distribution but absent from Africa. The name is from the Greek xylon, wood, and osma, odor, in an apparent reference to the fragrance of the wood of some species. In the Hawaiian Islands, there is 1 endemic species.

(SANDALWOOD FAMILY)

Parasitic and hemiparasitic shrubs, trees, and herbs. Key features are simple opposite leaves, which are reduced to scales in Exocarpus, the often thick and fleshy nature of the corolla, and the fleshy, drupaceous fruit. Recent molecular data have expanded the concept of the family by merging the Viscaceae, which includes the familiar Northern Hemisphere mistletoes, into the Santalaceae. A virtually cosmopolitan family with 44 genera and about 875 species worldwide, found primarily in the tropics and subtropical dry regions. Economically the family is important primarily for the production of sandalwood (Santalum), highly prized for its fragrant wood. Some species of sandalwood are cultivated. In the Hawaiian Islands represented by the indigenous genera Exocarpus, Korthalsella, and Santalum.

Zanthoxylum kauaense

Small to medium-sized trees 3–15 m tall, with trunks up to 40–50 cm in diameter. Leaves compound with 3–7 leaflets, leaflets thick, coriaceous, ovate to elliptic or suborbicular. Occurs in mesic forest and occasionally in dry or wet forest at elevations of 300–1,980 m on all of the main islands except Ni‘ihau and Kaho‘olawe. A highly variable plant, with many of the variants once recognized as species.

Santalaceae

Exocarpus—A genus of 26 species of

broom-form shrubs or sometimes small trees found in Southeast Asia, Malesia, Australia, and the Pacific, including the Hawaiian Islands. The name is from the Greek exo, out of, and karpos, fruit, referring to the drupe being partially embedded in a fleshy receptacle. In the Hawaiian Islands represented by 3 endemic species.

Exocarpus gaudichaudii A. DC.

au, heau, hulumoa, kaumahana ENDEMIC | RARE

4.514 

Xylosma hawaiiense—male plant

Trees up to 9 m tall. The leaves are variable in size and shape. Plants dioecious. The roundish fruit turns purple-black at maturity. Occurs in mesic forest and occasionally dry to wet forest at elevations of 400–1,220 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Unlike most of the other species of Xylosma elsewhere, the native Hawaiian species lack spines.

4.515 

Exocarpus gaudichaudii—habit

157

158

FLOWERING PLANTS

Stems cylindrical but abruptly flattened above the third node, 10–30 cm in length. Parasitic primarily on native woody plants in most vegetation types except coastal habitats and open bogs. In mesic and wet forest at elevations of 600–1,350 m or higher on all of the main islands except Ni‘ihau and Kaho‘olawe. The most common and widespread species. Outside the Hawaiian Islands, it is present on Henderson Island.

Stems cylindrical, then flattened after 1–2 internodes, 15–50 cm in length, branches many, oblique or decussate. Native to the Pacific Islands. Parasitic on native woody plants in diverse mesic forest at elevations of 300–1,220 m on Kaua‘i, O‘ahu, Lāna‘i (Kapua Gulch), and East Maui (Waikamoi Trail).

Korthalsella remyana Tiegh. hulumoa, kaumahana ENDEMIC | SECURE

Korthalsella latissima (Tiegh.) Danser ENDEMIC | SECURE

4.516 

Exocarpus gaudichaudii

Small broom-form trees or shrubs 2–5 m tall that are densely branched toward the tips. The leaves are scalelike and highly reduced, although sometimes a few foliaceous ones may be present. Uncommon on ridges in mesic forest and shrubland at elevations of 300–920 m or higher on all of the main islands except Kaua‘i.

Korthalsella—A genus of 25 species of mostly parasitic, brittle subshrubs or perennial herbs with strongly jointed stems that occur in eastern Africa, islands in the Indian Ocean, Asia, Japan, Malesia, Australia, New Zealand, and the Hawaiian Islands. The name honors P. W. Korthals, a nineteenth-century Dutch botanist. In the Hawaiian Islands represented by 4 endemic and 2 indigenous species. These taxa are thought to represent a single lineage.

4.520 

4.518 

Korthalsella latissima

Stems with 1–2 branches, these 10–20 cm in length. Branches conspicuously narrower than the main stem, distichous. Parasitic on native woody plants in wet and mesic forest at elevations of 1,100–1,220 m on Kaua‘i, O‘ahu (Mount Ka‘ala), and Hawai‘i (Waiākea Forest Reserve, Mauna Loa). Closely related to K. complanata.

Korthalsella complanata (Tiegh.) Engl.

Korthalsella platycaula (Tiegh.) Engl.

INDIGENOUS | SECURE

INDIGENOUS | SECURE

hulumoa, kaumahana

4.517 

Korthalsella complanata

hulumoa, kaumahana

4.519 

Korthalsella platycaula

Korthalsella remyana

Stems cylindrical or slightly flattened on the upper nodes, 10–15 cm in length, decussate. Parasitic on Metrosideros in dry to wet forest and open bogs at elevations of 300–1,220 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Santalum—A genus of shrubs and trees with fragrant flowers that are semiparasites on the roots of other plants. There are about 25 species found in eastern Malesia, Australia, Polynesia, and the Juan Fernandez Islands. The name is from the Greek santalon, sandalwood. In the Hawaiian Islands represented by 6 endemic species, 3 of which have redtinged flowers, 2 with fragrant greenish flowers, and 1 putative ancient hybrid between the clades (S. involutum). These species represent 2 separate founding lineages. A red-flowered Santalum lineage dispersed from Australia to the Hawaiian Islands approximately 1.0–1.5 million years ago and formed the S. freycinetianum clade. The second Santalum colonist arrived to the Islands approximately 0.4–0.6 million years ago and led to the white-flowered S. ellipticum/S. paniculatum clade. Sandalwood is highly prized for its fragrant wood.

FLOWERING PLANTS

Santalum ellipticum Gaudich.

‘iliahi, ‘iliahialo‘e, coast sandalwood ENDEMIC | SECURE

factor in altering the traditional Hawaiian way of life, where land and natural resources were not owned and barter was the only means of exchange.

Santalum haleakalae Hillebr. ‘iliahi

ENDEMIC | RARE/ENDANGERED

cones to upper-elevation wet forest at elevations of 450–2,000 m on Hawai‘i. The var. paniculatum is typically a shrub, while the pictured var. pilgeri is often a tree up to 20 m tall.

Santalum pyrularium A. Gray ‘iliahi

ENDEMIC | SECURE

4.521 

Santalum ellipticum

Sprawling shrubs or small trees 1–5 m tall with coriaceous or succulent leaves that are variable in shape and size. The fragrant greenish flowers are tinged with brown or orange. Occurs in dry shrub­ land and forest, often on a‘ā lava or rocky sites, and also along the coast on all of the main islands from sea level to 560 m elevation or more. Extinct on Kaho‘olawe and Laysan. A highly variable species in both vegetative and flowering characters.

Santalum freycinetianum Gaudich. ENDEMIC | SECURE

4.523 

Santalum haleakalae var. haleakalae

Small trees 2–4 m tall with stiff, coriaceous ovate leaves that are often purple tinged. The flowers open cream colored, turn dark red with age, and exhibit only a light fragrance. There are 2 named varieties. The rare var. haleakalae pictured here occurs in alpine shrubland at elevations of 1,900–2,700 m on dry slopes of Haleakalā, East Maui. The endangered var. lanaiense, formerly known as S. freycinetianum var. lanaiense, is found in dry forest and shrublands on slopes and ridges from 550–1,350 m elevation on Moloka‘i, Lāna‘i, and Maui.

Santalum paniculatum Hook. & Arn. ‘iliahi

ENDEMIC | SECURE

4.522 

Santalum freycinetianum

Small shrubs or trees 1–13 m tall with drooping branches and leaves that often appear wilted. Flowers turn dark red with age, weakly fragrant. Although 3 varieties were once recognized, the species is now considered to consist of a single taxon restricted in distribution to scattered locations at elevations of 300–800 m throughout the Wai‘anae and Ko‘olau mountains on O‘ahu. The sandalwood industry, which thrived on the Hawaiian Islands in the early 1800s, caused a great depletion of this species. Payment for harvesting rights was made in money and is thought to have been a significant

4.524 

Santalum paniculatum var. pilgeri

Shrubs or trees 3–20 m tall with yellowish orange to olive green leaves, glossy above and pale and dull beneath. The highly fragrant flowers are greenish in color when mature and tinged with brown or orange. Occurs in dry woodland on lava or cinder

4.525 

Santalum pyrularium

A red-flowered species formerly considered to be a variety of S. freycinetianum but now considered to be a distinct species. It differs in having more narrowly elliptic leaves that are fully expanded and not appearing wilted as in S. freycinetianum. Occurs in mesic forests from 270–1,440 m on Kaua‘i; common around the state parks of Waimea Canyon and Kōke‘e, where it prefers partial to full shade. Listed in the Manual as S. freycinetianum var. pyrularium.

Sapindaceae

(SOAPBERRY FAMILY)

Trees, shrubs, vines, lianas, or rarely herbs. Key features are the typically pinnate leaves, circinately coiled tendrils in the climbing genera, small flowers that are usually unisexual or in those that are bisexual functioning as unisexual, and the seeds generally with an aril. There are about 131 genera and about 1,450 species that are found predominately in the tropical and subtropical regions of the world. Economically the family is of little importance. Lychee (Litchi) is cultivated for its edible fruit. Cultivated ornamentals include the golden rain tree (Koelreuteria), tuckeroo (Cupaniopsis), soapberry (Sapindus), Mexican buckeye (Ungnadia), and balloon vine (Cardiospermum). In the Hawaiian Islands represented by 3 indigenous genera, Alectryon, Dodonaea, and Sapindus.

159

160

FLOWERING PLANTS

Alectryon—A genus of trees and some-

times shrubs with about 34 species from Australia, New Zealand, Malesia, and Polynesia. The name is from the Greek alektryon, cock, referring to the red aril that covers the seed coat resembling a cock’s comb. In the Hawaiian Islands represented by 1 endemic species with 2 distinct varieties.

Alectryon macrococcus Radlk. ‘ala‘alahua, māhoe

ENDEMIC | ENDANGERED

Dodonaea—A pantropical and warm tem-

perate genus of shrubs or small trees with either simple or compound leaves, with a distribution centered in Australia. There are 68 species, one of which is pantropical. The name honors Rembert Dodoens, a sixteenth-century court physician and professor of medicine at Leiden. In the Hawaiian Islands represented by 1 indigenous, highly variable species. At least 4 other species were recognized from Hawai‘i in the past based on the morphology of the fruit.

medicine to reduce inflammation and cleanse wounds. In the Hawaiian Islands represented by 1 endemic species with simple leaves and 1 widely dispersed species with compound leaves. These species represent separate founding lineages.

Sapindus oahuensis Hillebr. ex Radlk. āulu, kaulu, lonomea, soapberry ENDEMIC | SECURE

Dodonaea viscosa Jacq. ‘a‘ali‘i, kūmakani

INDIGENOUS | SECURE

4.529 

4.526 

Evergreen trees 6–15 m tall with simple ovate to elliptic leaves. Occurs in mesic to dry forest at elevations of 60–610 m on Kaua‘i and in both mountain ranges on O‘ahu. The seeds were sometimes strung into lei.

Alectryon macrococcus var. auwahiensis—

habit

4.528 

4.527 

Alectryon macrococcus var. macrococcus

Trees 4–11 m tall with reddish brown branches and pinnately compound leaves. The brown, glossy seeds are covered by a scarlet aril. Occurs in dry to mesic forest at elevations of 360–1,070 m. Two varieties are recognized. The var. auwahiensis has the lower leaf surfaces densely pubescent and occurs only on East Maui. When censused in 2002, this variety had only 2 small extant populations with 22 individuals. The fleshy arils and seeds were eaten in the past. This plant is declining because insects and rats destroy most of the seeds. The var. micrococcus, with the lower leaf surface sparsely pubescent, occurs in dry to mesic forest on Kaua‘i, O‘ahu, Moloka‘i, and West Maui. There are currently 15 populations and about 370 extant trees in the Wai‘anae Mountains.

Sapindus oahuensis

Dodonaea viscosa

Shrubs or small trees 2–8 m tall, pantropical in distribution. Leaves variable in shape, ranging from elliptic to lanceolate, and 3–10 cm or more in length. The species is usually dioecious. The fruit varies in color from straw colored to dark reddish brown. In the Hawaiian Islands occasionally dominant in a wide variety of habitats in dry, mesic, and wet forest and in subalpine shrubland at elevations ranging from sea level to 2,350 m on all of the main islands. A very polymorphic plant, with many of the variants previously recognized as species. The fruit and leaves are used in making lei. A red dye for kapa was made from the fruit. The wood was also utilized for building houses or fashioned into spears.

Sapindus—A pantropical genus of trees consisting of 10 species, with diversity centered in Southeast Asia and Malesia. The name is from the Latin sapo, soap, and indicus, Indian, “Indian soap” referring to the saponin found in the fruit. This chemical lathers in water and was used as a soap substitute as well as in traditional

Sapindus saponaria L. a‘e, mānele

INDIGENOUS | SECURE

4.530 

Sapindus saponaria

Deciduous trees to 25 m tall, with the bark on mature plants falling off in large scales. The leaves are evenly pinnately compound. Distributed from Mexico and South America west across the Pacific Basin to New Caledonia and also present in Africa. Occurs in mesic forest at elevations of 900–1,370 m on Mauna Loa and Hualālai, Hawai‘i. These plants are somewhat different from populations outside of Hawai‘i. The seeds were strung into lei, necklaces, and rosaries.

FLOWERING PLANTS

Sapotaceae

(SAPODILLA FAMILY)

Trees and shrubs. Key features are the presence of ducts or sacs that produce a milky latex, a completely septate superior ovary, and the fruit a berry with a thin leathery or bony outer covering. There are 53 genera and about 975 species, with a pantropical distribution for the family. Economically the family is important for chewing gum from the chicle plant (Achras); gutta-percha of commerce, which comes from the milky sap of several genera; and edible fruits such as the sapodilla (Achras); sapote (Calocarpum); eggfruit (Pouteria); and the star apple (Chrysophyllum). In the Hawaiian Islands represented by the indigenous genera Planchonella and Sideroxylon.

Planchonella—A tropical and subtropical

genus of trees and shrubs with about 100 species. Flowers bisexual or seldom unisexual, solitary or several in axillary clusters. Distributed from India through Asia to Australia and the Pacific, with 2 species in South America. Named in honor of Jules Émile Planchon. Currently represented in the Hawaiian Islands by 1 variable species in need of further study.

forest at elevations of 240–980 m on all of the main islands except Ni‘ihau and Kaho‘olawe. A highly variable plant, with many of the forms given species status in the past. The milky sap was used as birdlime to catch small birds. The hard wood was used in house building and for digging sticks and spears. Listed in the Manual as Pouteria sandwicensis.

Sideroxylon—A genus of about 70 species

of tropical and subtropical trees characterized by gummy or milky sap and extremely hard wood. Largely neotropical in distribution but present across the Pacific to Madagascar and Africa. The name is derived from the Greek words sideros, iron, and xylon, wood. In the Hawaiian Islands represented by 1 indigenous species.

Sideroxylon polynesicum (Hillebr.) Smedmark & Anderb. keahi INDIGENOUS | VULNERABLE

Scrophulariaceae (FIGWORT FAMILY)

Annuals, herbaceous perennials, shrubs, and trees. Widely distributed in the tropics and warm temperate areas of the world. The family has been quite dramatically redefined on the basis of molecular data, and many genera once placed here have now been moved to other families (e.g., Orobanchaceae, Phrymaceae, Plantaginaceae, and Stilbaceae), while the Myoporaceae and Buddlejaceae have been merged into the Scrophulariaceae. As now defined, the family has 54 genera and about 1,800 species. It includes a great many species cultivated as ornamentals, including Buddleja, Eremophila, Myoporum, Verbascum, and Sutera. In the Hawaiian Islands represented by the indigenous genus Myoporum.

Bacopa— Once placed in the Scrophulariaceae, Bacopa is now considered to be placed within the Plantaginaceae. Myoporum—A genus of shrubs and small

trees with 32 species in Australia, New Guinea, New Zealand, Mauritius, and the Pacific Islands. The name is from the Greek myo, close, and poros, pore, as the leaves are often glandular-punctate. This genus was formerly placed in the Myorporaceae. The Hawaiian Islands have 1 endemic and 1 indigenous species found in a wide variety of habitats.

Planchonella sandwicensis (A. Gray) Pierre ‘āla‘a, āulu, kaulu

ENDEMIC | SECURE

Myoporum sandwicense A. Gray naio, bastard sandalwood INDIGENOUS | SECURE

4.532 

4.531 

Planchonella sandwicensis

Shrubs or trees to 12 m tall with white latex and variable-shaped leaves. The fruit is yellow, orange, or purplish black at maturity. Occurs in dry to mesic

Sideroxylon polynesicum

Shrubs or trees to 10 m tall with white latex and coriaceous leaves. Dioecious, with scented flowers present on separate plants; male flowers with 7–10 stamens. Found in the Austral Islands, Rapa Nui, and the Hawaiian Islands, where it is uncommon on slopes and remnant patches of dry forest at elevations of 130– 640 m on all of the main islands except Ni‘ihau, Kaho‘olawe, and Hawai‘i. Listed in the Manual as Nesoluma polynesicum.

4.533 

Myoporum sandwicense

161

162

FLOWERING PLANTS

Shrubs to small trees 1–10 m tall, with many branches. The leaves are somewhat fleshy and variable in shape. Flower color varies from pure white to white with purplish blotches or even pink. Native outside of the Hawaiian Islands in the Cook Islands. Occurs in strand vegetation, dry forest, lava fields, and mesic to wet forest and often becomes a codominant in subalpine forest on all of the main islands except Kaho‘olawe at elevations from sea level to 2,380 m. A highly variable plant, both morphologically and ecologically. The wood when burned has a smell similar to sandalwood. It was formerly used for house frames.

Solanaceae

(POTATO FAMILY)

Herbs, shrubs, trees, and lianas. Key features are alternate simple leaves and an absence of stipules. The sympetalous flowers are bisexual and actinomorphic or only slightly zygomorphic. Fruit typically a berry but occasionally a capsule. A widely distributed family with a center of diversity in the neotropics. There are 91 genera and about 2,450 species. Economically the family is important for food plants such as potatoes (Solanum), eggplant and tomatoes (Solanum), and peppers (Capsicum); for drug and medicinal plants such as tobacco (Nicotiana), henbane (Hyoscyamus), belladonna (Atropa), and stramonium (Datura); and for ornamentals such as petunia (Petunia), boxthorn (Lycium), cup of gold (Solandra), angel’s trumpet (Brugmansia), and nightshade (Solanum). In the Hawaiian Islands represented by 1 endemic genus, Nothocestrum, and 2 indigenous genera, Lycium and Solanum. Species of Brugmansia, Capsicum, Cestrum, Datura, Lycopersicon, Nicandra, and Nicotiana have naturalized in the Islands.

Lycium—A genus of about 80 species of woody shrubs, usually spiny, widely scattered in temperate and tropical America, Australia, Eurasia, and the Pacific Islands, with a center of diversity in tropical America. The name is from the Greek lykion, which is the vernacular name for a shrub from southwestern Turkey. In the Hawaiian Islands represented by 1 indigenous species.

Lycium sandwicense A. Gray

‘ae‘ae, ‘ākulikuli ‘ae‘ae, ‘akulikuli ‘ōhelo, ‘ākulikuli kai, ‘ōhelo kai INDIGENOUS | SECURE

Trees 10–12 m tall with a thick trunk up to 45 cm diameter. Leaves oblong and clustered near the ends of the branches, relatively small at 5–12 cm in length. Flowers numerous on short axillary spurs. The red-orange fruit is enclosed by the persisting calyx. Rare in dry to mesic forest at elevations of 580–1,830 m on Hawai‘i (Ka‘ū District north to Waimea) (see figure 8.3). There are only 6 remaining populations with about 50 individuals in total. Threats have come from habitat loss, feral grazing animals, alien plant species, and fire.

Nothocestrum latifolium A. Gray ‘aiea, hālena 4.534 

Lycium sandwicense

Spreading shrubs to 1 m tall with stiff branches and somewhat succulent leaves clustered on condensed lateral shoots. Fruit, a 2-celled berry, is succulent and red at maturity. Indigenous to Rapa Nui, Tonga, Juan Fernandez Islands, and the Hawaiian Islands, where it is found in subsaline and rocky coastal sites from sea level to elevations of 40 m on all of the main islands. The fruit is edible but slightly salty.

Nothocestrum—An

endemic Hawaiian genus of 4 species of softwood trees that are uncommon or rare. The name is from the Greek nothos, false, and Cestrum, a genus of American shrubs. The gnarled, often scraggly appearance of these plants is distinctive. The genus is the host plant for the endangered Blackburn’s sphinx moth (Manduca blackburni), one of the Hawaiian Islands’ largest native insects with a wingspan of up to 12 cm.

Nothocestrum breviflorum A. Gray

‘aiea, hālena

ENDEMIC | RARE

4.536 

Nothocestrum latifolium

Small trees to 10 m tall with a gnarled trunk. Leaves usually deciduous during the dry season, thick, broadly ovate to suborbicular and usually lobed, 4–20 cm in length. The fragrant flowers are several to numerous in clusters on short spurs. Occurs in dry to mesic forests at elevations of 460–1,530 m on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui.

Nothocestrum longifolium A. Gray ‘aiea, hālena

ENDEMIC | SECURE

ENDEMIC | ENDANGERED

4.535 

Nothocestrum breviflorum

4.537 

Nothocestrum longifolium

FLOWERING PLANTS

Shrubs or small trees to 6 m tall. Leaves thin and lanceolate to elliptic–oblong, 10–32 cm in length, widely spaced but sometimes also clustered at the ends of the branches. Occurs primarily in wet forests but occasionally in mesic forests at elevations of 360–1,620 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Solanum nelsonii Dunal

‘ākia, pōpolo, nightshade ENDEMIC | VULNERABLE

Sterculiaceae

(CACAO

FAMILY)—

Molecular data have now suggested that the Sterculiaceae, along with the Bombacaceae and Tiliaceae, should now be merged into the Malvaceae. This includes Waltheria, the 1 Hawaiian genus in the former Sterculiaceae.

Ternstroemiaceae—A name formerly used for what is now the Pentaphylacaceae.

Nothocestrum peltatum Skottsb. ‘aiea, hālena

ENDEMIC | ENDANGERED

4.539 

Solanum nelsonii

Sprawling or trailing shrubs to 1 m tall forming large clumps. The ovate to rounded leaves are covered with stellate hairs. Common in coastal sites in coral rubble or sand from sea level to elevations of 150 m on Kure, Midway (extinct), and Pearl and Hermes atolls, Laysan (extinct), Nīhoa, and Ni‘ihau, but it is thought to be extinct on Kaua‘i and O‘ahu and is relatively rare on Moloka‘i, Maui, and Hawai‘i.

Solanum sandwicense Hook. & Arn. 4.538 

Nothocestrum peltatum

pōpolo ‘aiakeakua, nightshade ENDEMIC | ENDANGERED

Small trees to 5 m tall with coriaceous ovate leaves that are typically peltate. A rare plant from diverse mesic forest at elevations of 915–1,220 m on Kaua‘i (Kalalau Lookout, Kumuwela, Nu‘alolo, Mākaha Valley, west side of Waimea drainage). The most recent estimates are that there are 7 extant populations totaling only 23 individuals. The most serious threat has come from competition with alien plant species, with fire also of concern.

Solanum—A large genus of herbs, shrubs,

trees, lianas, or vines with about 1,250 species that are cosmopolitan in distribution. The name is from the Latin used to designate a member of this genus. A highly diversified group, many of which are important as food plants and ornamentals. In the Hawaiian Islands, there are 3 endemic and 1 indigenous species and 7 species that have naturalized. The 3 endemic species represent a single colonizing lineage separate from that of the indigenous species. Some of the introduced species have become invasive weeds.

4.540 

Solanum sandwicense

Large sprawling shrubs to 4 m tall, with all parts of the plant covered with dense yellowish brown stellate hairs. Rare in diverse mesic forest at elevations of 760– 1,220 m on Kaua‘i and O‘ahu (extinct). Known today from 5 populations with a total of only 18–20 individuals on Kaua‘i. The last plant on O‘ahu died in the 1990s, but seeds from this individual, saved before its death, have now been grown and planted out in the field. Threats have come from feral grazing animals, alien plant species, and rats.

Theaceae (TEA FAMILY)—The genus Eurya in the Hawaiian flora was formerly considered as a member of the Theaceae but is now placed in the Pentaphylacaceae. Thymeleaceae (DAPHNE FAMILY)

Shrubs and trees. Key features are the presence of stems and branches with strong and tough fibers and an unusual flower structure with sepals, petals, and stamens difficult to separate. The floral tube appears to be a calyx or corolla but is actually a hollow receptacle. What appear to be petals are actually stipular appendages of the sepals. The flowers are typically fragrant at night. There are 45 genera and about 850 species that are found around the world in the tropics, subtropics, and temperate regions. Economically the family is of little importance except for a few cultivated ornamentals, such as leatherwood (Dirca), paper bush (Edgeworthia), rice flower (Pimelea), mezereum (Daphne), and Dais. In the Hawaiian Islands represented by the indigenous genus Wikstroemia.

Wikstroemia—A genus of shrubs and sub-

shrubs and sometimes small trees, with about 70 species found in Southeast Asia, Australia, Fiji, and the Hawaiian Islands. The name honors J. E. Wikström, a nineteenth-century Swedish botanist. In the Hawaiian Islands, there are 12 recognized endemic species, which are often difficult to separate, representing a single colonizing lineage. Flowers in the genus are typically perfect, but in Hawaiian species they may be perfect, unisexual (dioecious), or with a mix of perfect and unisexual flowers (polygamous). The fibers of the plants are extremely strong, making it very difficult to break off an individual branch.

163

164

FLOWERING PLANTS

flowering rachis elongates up to 50 mm in fruit. Scattered in occurrence in Pandanus forest and a common species in mesic forest at elevations of 380–1,220 m on Kaua‘i.

These fibers were traditionally used for making cordage. Poisonous alkaloids are common in the genus, and Native Hawaiians used powdered roots to immobilize fish near the shore and in fishponds to make them easier to catch.

Wikstroemia oahuensis (A. Gray) Rock

Wikstroemia bicornuta Hillebr.

ENDEMIC | SECURE

‘ākia

‘ākia

with silky pubescence or glabrous. Occasionally a dominant species in open dry forest or shrubland, a‘ā lava fields, or even mesic forest at a wide range of elevations from sea level to 2,290 m on Hawai‘i.

Wikstroemia pulcherrima Skottsb. ‘ākia

ENDEMIC | SECURE

ENDEMIC | RARE

4.543  4.541 

Wikstroemia bicornuta

Straggling shrubs to small trees 1.0–2.5 m or more tall, stems pubescent to glabrous. Leaves 3.5–8.0 cm in length, dark green above, elliptic to suborbicular, apex and base obtuse to broadly rounded, chartaceous. Flowers unisexual, greenish, pubescent. Occurs in wet forests at elevations of 900–1,059 m on Lāna‘i and West Maui (Mount ‘Eke).

Wikstroemia furcata (Hillebr.) Rock ‘ākia

ENDEMIC | SECURE

Wikstroemia oahuensis var. palustris

Sparsely branched shrubs or small trees to 4 m tall, with many small twigs. Leaves 2.5–8.0 cm in length, elliptic to slightly ovate, thin and transparent to firmly coriaceous, lower surface with conspicuous lateral veins, apex and base acute or rounded. Flowers perfect or unisexual, green to yellow, pubescent to glabrous. Two varieties are recognized, with var. oahuensis common in mesic to wet forest and wet ridges and bogs at elevations of 100–1,400 m on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui. The bog var. palustris, pictured, occurs only on Kaua‘i and is a densely branched, low shrub. A highly variable species.

Wikstroemia phillyreifolia A. Gray ‘ākia, kahui

ENDEMIC | SECURE

4.545 

Wikstroemia pulcherrima

Low shrubs or small, densely branched trees up to 2 m tall, young branches pubescent or with scattered hairs. Leaves 2–3 cm in length, soft coriaceous, elliptic to ovate, apex and base acute to rounded. Flowers perfect or unisexual and plants then dioecious, yellow, with long silky hairs. Common in grazed pastures, roadsides, and remnant dry forests at elevations of 700–950 m on Hawai‘i (South Kohala and North Kona districts).

Wikstroemia uva-ursi A. Gray ‘ākia, kauhi

ENDEMIC | SECURE

4.544  4.542 

Wikstroemia furcata

Shrubs or small trees to 4.5 m tall, stems typically with a mat of rust-colored hairs. Leaves 7–12 cm in length, dark green above, elliptic to ovate, apex acute to rounded. Flowers perfect, greenish to yellowish green, usually pubescent. The

Wikstroemia phillyreifolia

Low, sprawling shrubs or small bushy trees to 3 m tall, often with a single stem, young branches pubescent or glabrous. Leaves 2–4 cm in length, coriaceous, elliptic to suborbicular, apex and base acute or rounded. Flowers both perfect and unisexual and plants thus polygamous, yellow,

4.546 

Wikstroemia uva-ursi var. uva-ursi

Densely branching, sprawling, or prostrate shrubs to 1.5 m tall, young stems

FLOWERING PLANTS

with dense yellow to reddish brown pubescence. Leaves thick and firm, 1.7– 3.0 cm in length, elliptic to orbicular. Flowers typically unisexual. Infrequent on open hillsides, dry ridges, a‘ā lava, and other rocky sites in coastal areas at low elevations up to 420 m or slightly higher. There are 2 named varieties, with the var. kauaiensis restricted to Kaua‘i and the pictured var. uva-ursi on O‘ahu, Moloka‘i, and Maui. Often cultivated because of its attractive foliage and red fruit.

Boehmeria grandis (Hook. & Arn.) A. Heller ‘ākōlea, false nettle ENDEMIC | SECURE

Urticaceae

(NETTLE FAMILY)

Herbs, shrubs, or rarely trees or lianas. Key features are the usual presence of stinging hairs on the stems and leaves, a simple ovary with a single style, and a cymose inflorescence borne on short axillary shoots. There are 55 genera and about 1,650 species that are primarily found in the tropics and subtropics but also distributed in temperate regions. The name comes from the Latin word urtica, for nettle, which is derived from the verb urere, to burn. Hawaiian members of the family are unusual in that they do not have stinging hairs, with the exception of Hesperocnide, which is thought to be a recent offshoot from its California relative. The loss of the trait is hypothetically due to the absence of mammalian herbivores. Economically the family is of minor importance. Ramie, a commercial fiber, comes from Boehmeria. A few species of Pilea and Soleirolia are grown for ornamental value. In the Hawaiian Islands represented by 2 endemic genera, Neraudia and Touchardia, and 5 indigenous genera: Boehmeria, Hesperocnide, Pilea, Pipturus, and Urera. One weedy species of Urtica and 1 species of Pilea have naturalized. Boehmeria—A pantropical genus of about 80 species of shrubs or small trees found with some extension into north temperate areas. Stems with watery sap. Leaves alternate or opposite but opposite in the Hawaiian species, typically with 3 main veins. The name honors Georg Rudolph Boehmer, an eighteenth-century German botanist. In the Hawaiian Islands, there is 1 endemic species. Boehmeria nivea, the source of ramie fiber, is still grown in the Islands but not on a commercial basis.

4.547 

Boehmeria grandis

Shrubs 1–2 m tall with thick, coriaceous, ovate to elliptic leaves. Unique among the native Hawaiian Urticaceae in having opposite leaves. Monoecious, with male and female flowers separate and the males concentrated toward the ends of the flowering branches. Occurs along streambeds, on slopes, ridges, and valleys in mesic to wet forest at elevations of 150– 1,100 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Erect annual herbs to 60 cm tall with stinging hairs on all parts of the plant. Locally common in subalpine woodland or alpine areas at elevations of 1,780– 2,400 m on the saddle between Hualālai, Mauna Loa, and Mauna Kea, Hawai‘i. Closely resembles the introduced stinging nettle, Urtica urens.

Neraudia—An endemic Hawaiian genus of shrubs, sometimes climbing, or small trees, with 5 closely related dioecious species with milky sap representing a single colonizing lineage. Leaves alternate and with 3 main veins. Fruiting calyx fleshy, fruit not embedded in a fleshy receptacle. The name honors J. Néraud, a nineteenth-century amateur botanist of Madagascar. The plants were of importance to the Hawaiians because of the sturdy fibers used in the making of kapa.

Neraudia kauaiensis (Hillebr.) R. S. Cowan ‘oloa, ma‘aloa

ENDEMIC | RARE

Hesperocnide—A genus of 2 species of erect annual herbs, with 1 species endemic to California and the other to the island of Hawai‘i. The name is from the Greek hespera, west, and knide, nettle, referring to the geographic distribution of a species of the plant. It closely resembles the common nettle, Urtica urens, which has naturalized in the Hawaiian Islands.

Hesperocnide sandwicensis (Wedd.) Wedd. ENDEMIC | SECURE

4.549 

4.548 

Hesperocnide sandwicensis

Neraudia kauaiensis

Erect shrubs 1.5–3.0 m tall with branches pubescent with erect and ± appressed hairs. Lower leaf surface moderately pubescent with white or gray hairs. Upper leaf surface with a conspicuous tuft of hairs at its base. Occurs in diverse mesic forest at elevations of 620–1,070 m on Kaua‘i.

165

166

FLOWERING PLANTS

Neraudia melastomifolia Gaudich. ‘oloa, ma‘aloa

ENDEMIC | SECURE

Low, short-lived perennial herbs with fleshy stems and ovate or rounded leaves. Monoecious, with male and female flowers on the same plant. Found in northern South America, Galapagos Islands, and the Hawaiian Islands, where it occurs in streambeds, moist cliffs, and seeps in mesic and wet forest at elevations of 40–1,950 m on all of the main islands and on Lehua.

Pipturus—A genus of shrubs or small trees,

4.550 

Neraudia melastomifolia

Shrubs or small trees up to 4 m tall with thick or thin leaves and milky sap. Lower leaf surface glabrous or sparsely pubescent along the veins. Occurs in mesic forest and diverse mesic forest at elevations of 270–1,160 m on Kaua‘i, O‘ahu, Moloka‘i (Wāwā‘ia Gulch), and West Maui (Olowalu, ‘Īao Valley). A variable species.

with about 30 species found in Malesia, Australia, the Mascarene Islands, and several Pacific islands. Stems with watery sap. Leaves thin and alternate, with 3–5 main veins. Fruiting calyx dry, fruit embedded in a fleshy receptacle. Plants dioecious, monoecious, or gynodioecious. The name is from the Greek pipto, to fall, and oura, tail, referring to the stigma, which falls away. Represented in the Hawaiian Islands by 4 endemic species representing a single colonizing lineage. Another important plant to the Native Hawaiians, who used the fibers for cordage and in the making of kapa.

māmaki

ENDEMIC | SECURE

māmaki

ENDEMIC | SECURE

4.553 

Pipturus kauaiensis

Shrubs 1.5–3.0 m tall with slender branches. Leaves 6–18 cm in length, thin, ovate to broadly lanceolate, covered with fine white hairs on the lower surface. Plants gynodioecious. Occurs in mesic valleys and diverse mesic forest at elevations of 60–1,280 m on Kaua‘i.

Pilea peploides (Gaudich.) Hook. & Arn. INDIGENOUS | SECURE

Pilea peploides

Pipturus kauaiensis A. Heller

Pipturus albidus (Hook. & Arn.) A. Gray

Pilea—A large genus of herbs and semiwoody shrubs represented by about 650 species in the tropics and subtropics, especially South America but missing from Australia. The name is from the Latin pileus, a felt cap, referring to the shape of the large pistillate calyx lobe of the type species. In the Hawaiian Islands represented by 1 indigenous species and the naturalized artillery plant, Pilea microphylla.

4.551 

Shrubs or small trees 2–6 m tall with relatively thick branches. Leaves 7–20 cm in length, highly variable in shape, texture, and color. Plants dioecious or monoecious. Occurs in mesic valleys, mesic to wet forest, and diverse mesic forest at elevations of 70–1,870 m on all of the main islands except Ni‘ihau and Kaho‘olawe. A polymorphic plant throughout its range. The plant is the host for the Kamehameha butterfly (Vanessa tameamea), one of only two species of butterfly native to the Hawaiian Islands. The fibers of the inner bark were used for making kapa and for cordage. The waxy-white fruit was formerly used as a laxative and as a remedy for thrush in children.

4.552 

Pipturus albidus

Touchardia—An endemic Hawaiian genus of shrubs with 1 variable species. Stems with viscid sap. Leaves alternate with 3 main veins. Flowers in dense globose panicles. The name perhaps honors a Mr. Touchard, but the derivation is uncertain. This plant was also highly valued by the Hawaiians for its strong fibers, which were utilized in a variety of ways.

FLOWERING PLANTS

Touchardia latifolia Gaudich. olonā

ENDEMIC | SECURE

passed down between generations. The strength and durability of Touchardia ropes made them highly prized for European mountain climbers in the nineteenth century, with even short sections of rope selling for large sums.

Urera kaalae Wawra ōpuhe

ENDEMIC | ENDANGERED

Urera—A pantropical genus of shrubs or

small trees with about 35 species that occur in tropical Africa, the Americas, Madagascar, and the Pacific Islands. Most species have stinging hairs, but these are absent in the Hawaiian plants. Leaves alternately with 3–5 main veins or pinnately veined, margins entire or lobed. Inflorescence an open panicle. Plants dioecious or monoecious. The name is from the Latin uro, sting, referring to the presence of stinging hairs in many species outside of the Hawaiian Islands. In the Hawaiian Islands represented by 2 endemic species, which are thought to represent separate colonizing lineages.

Urera glabra (Hook. & Arn.) Wedd. hona, hōpue, ōpuhe ENDEMIC | SECURE

4.554 

Touchardia latifolia—habit

4.557 

Urera kaalae

Small trees or shrubs 3–7 m tall lacking stinging hairs. The pale green, thin, heartshaped leaves are somewhat asymmetrical. A rare plant from gulches in diverse mesic forest at elevations of 300–760 m in the Wai‘anae Mountains (Nāpepeiauolelo Gulch, ‘Ēkahanui near Pu‘ukaua, Kalua‘a near Pu‘uhāpapa), O‘ahu. The most recent surveys have found 4 populations with a total about 40 individuals. Threats have come from feral pigs and alien plant species.

Verbenaceae

(VERBENA FAMILY)—

Many genera of the Verbenaceae, including the genus Vitex present in the Hawaiian Islands, have now been merged into the Lamiaceae.

Violaceae

(VIOLET FAMILY)

4.555 

Touchardia latifolia

Shrubs 1–3 m tall, sparsely branched, with dark green, ovate to lanceolate leaves and unisexual flowers in dense globose clusters. Occurs in mesic valleys and wet forest at elevations of 70–1,200 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Touchardia produces the strongest natural fiber and thus was an important plant for the Hawaiians, who used the fibers for fishnets, rope for hauling and lashing canoes, and as a base for raincoats and feather capes that were

4.556 

Urera glabra

Shrubs or small trees 2–8 m or more in height and diameters up to as much as 30 cm. Foliage lacks stinging hairs. Leaves dark green with 3 main veins and multiple pairs of major pinnate veins. Plants dioecious. Occurs in mesic to wet forest and diverse mesic forest at elevations of 150–1,700 m on all of the main islands except Ni‘ihau and Kaho‘olawe. A highly variable plant. Utilized by the Hawaiians for its strong fibers to make fishnets and sometimes kapa cloth.

Herbs, shrubs, or rarely small trees or lianas. Key features are the 5-merous flowers, stamen joined in a ring, corolla often zygomorphic. There are 24 genera and about 700 species distributed widely in both tropical and temperate regions. Economically the family is of minor importance. Over 100 species of violets (Viola) are cultivated as ornamentals. In the Hawaiian Islands represented by 1 endemic genus, Isodendrion, and the indigenous Viola.

Isodendrion—An endemic Hawaiian genus of 4 species of shrubs, all rare. The name is from the Greek isos, equal, and dendron, tree, referring to the almost equal petals and the woody nature of the plants. Flowers solitary on short peduncles, with

167

168

FLOWERING PLANTS

pedicels less than 8 mm in length. One species is currently considered extinct in the wild.

Isodendrion laurifolium A. Gray aupaka

ENDEMIC | ENDANGERED

Isodendrion hosakae H. St. John aupaka

ENDEMIC | ENDANGERED

4.560 

4.558 

Isodendrion hosakae—habit

Isodendrion laurifolium

Wandlike small shrub with a few erect branches 1–2 m in length, leafy for 10–30 cm below the top. Leaves somewhat coriaceous, oblong-elliptic to narrowly elliptic, 4–16 cm in length, glabrous, veins conspiculously reticulate and raised on both surfaces, margins undulate. Petals purple with greenish white margins. Occurs in diverse mesic forest at elevations of 550–610 m on Kaua‘i and both mountain ranges on O‘ahu. Population estimates are less than 1,000 extant individuals, with most of these on Kaua‘i. There are just 5 populations with 24–64 individu­ als on O‘ahu. Threats come from feral pigs and goats and alien plant species.

Isodendrion longifolium A. Gray aupaka

ENDEMIC | VULNERABLE

4.559 

Wandlike, small shrub with a few erect branches 60–200 cm in length, leafy for 10–20 cm below the top. Leaves somewhat coriaceous, oblanceolate, 10–30 cm in length, glabrous, margins flat or only slightly undulate. Petals purple. Uncommon to rare on steep slopes and cliffs in diverse mesic and wet forest at elevations of 240–700 m on Kaua‘i (Hanapēpē, Wahiawa Mountains, Hā‘upu, and Limahuli, Wainiha, and Hanakāpī‘ai valleys) and the Wai‘anae Mountains of O‘ahu (Palikea, Pu‘upane). Population estimates are less than 1,000 extant individuals, with most of these on Kaua‘i. There are just 4 populations with 32–36 individuals on O‘ahu. Threats come from feral pigs and goats and alien plant species.

Viola—A large genus of herbs, in the

Hawaiian Islands becoming shrubby. Approximately 400 species widely distributed, especially in temperate regions of the world, with a center of biodiversity in the Andes. Distinguished from Isodendrion by petals conspiculusly uneven in size and sepals prolonged at their base. Flowers 1–2(5) on long peduncles and pedicels more than 10 mm in length. The name is from the Latin viola, a reference to a number of fragrant flowers. In the Hawaiian Islands represented by 7 endemic species of mostly shrubs or subshrubs, representing a single colonizing lineage. In some species, morphologically distinct cleistogamous flowers are produced.

Viola chamissoniana Ging. pāmakani, ‘olopū

ENDEMIC | ENDANGERED/SECURE

Isodendrion hosakae—fruit

Branched shrubs 30–80 cm tall, leafy on the upper 20 cm of the branches, stems pubescent. Leaves coriaceous, narrowly elliptic, 3–7 cm in length, veins impressed on upper surface and raised on lower surface, pubescent at least along the veins. Petals greenish yellow. A rare plant known from dry shrubland at elevations of 900–1,030 m on three cinder cones in the South Kohala District on Hawai‘i. Known from 3 populations and somewhere in the range of 320–1,000 indiviuals. Invasive grass species and associated fire have caused a sharp population decline.

4.561 

Isodendrion longifolium—habit

4.562 

Viola chamissoniana subsp. robusta

FLOWERING PLANTS

Small, erect, unbranched shrubs 30–80 cm tall. Leaves clustered on the upper part of the stem, lanceolate to elliptic lanceolate, 7.5–13.0 cm in length, glabrous. Flowers apparently all chasmogamous, petals 1–2 in upper leaf axils, petals pale lavender or white. Extremely rare and known only from a few extant plants at about 610 m elevation along Wahiawa Stream on Kaua‘i. Threats come from feral pigs and alien plant species.

Viola maviensis H. Mann ENDEMIC | SECURE

Viola kauaensis A. Gray

nani Wai‘ale‘ale, pohe hiwa ENDEMIC | SECURE

4.563 

Viola chamissoniana subsp. tracheliifolia

Slender, few-branched, or unbranched shrubs up to 1 m tall, prominent taproot. Leaves clustered on the upper part of the branches, deltate ovate to rarely cordate, 2.5–9.0 cm in length, glabrous or lower surface pubescent. Flowers apparently all chasmogamous, 1–2(4) per peduncle, petals white tinged purple at their base. Occurs in mesic to wet forest at elevations of 330–1,890 m on Kaua‘i, O‘ahu, Moloka‘i, and Maui. The subsp. chamissoniana is an endangered plant from dry cliffs in the Wai‘anae Mountains. Not threatened are the subsp. robusta from wet forests on Moloka‘i and subsp. tracheliifolia from mesic forests on Kaua‘i, O‘ahu, Moloka‘i, and Maui. This is the world’s tallest species of Viola.

Viola helenae C. N. Forbes & Lydgate ENDEMIC | ENDANGERED

4.564 

Viola helenae

4.565 

Viola kauaensis var. kauaensis

Perennial herbs with ascending to decumbent stems 10–50 cm in length. Leaves widely spaced, heart shaped, 2–5 cm in length, 3.5–6.0 cm in width. Flowers solitary in the leaf axils, petals white, lower surface pale blue to white, both cleistogamous and chasmogamous flowers present. Occurs in open bogs at 630–640 m elevation in Wahiawa Bog and the Alaka‘i Swamp at 1,150–1,175 m, Kaua‘i, and a boggy area of the Ko‘olau Mountains on O‘ahu at an elevation of about 630 m. The var. kauaensis occurs on Kaua‘i (Waineki to Mount Wai‘ale‘ale) and in the Ko‘olau Mountains on O‘ahu (between Kawainui and Kaipapa‘u). The var. wahiawaensis is an endangered taxon from Wahiawa Bog and Mount Wai‘ale‘ale.

4.566 

Viola maviensis—habit

4.567 

Viola maviensis

Dwarf subshrubs with erect to ascending stems 30–80 cm in length, short leafy segments of the stems alternating with long rhizomatous internodes. Leaves broadly ovate to rarely subcordate, 1.5–9.0 cm in length, margins with long, stiff teeth. Flowers fragrant, apparently all chasmogamous, petals dark violet to pale blue in color and usually arrayed in pairs per peduncle. Occurs in open bogs or sometimes bog margins, often in tussock vegetation, at elevations of 1,220–2,010 m on Moloka‘i (Pēpē‘ōpae Bog), Maui, and the Kohala Mountains of Hawai‘i. It hybridizes with Viola chamissoniana subsp. robusta on Moloka‘i where the two grow together.

169

170

FLOWERING PLANTS

Viola wailenalenae (Rock) Skottsb. ENDEMIC | SECURE

Zygophyllum are grown for ornament. In the Hawaiian Islands represented by the indigenous genus Tribulus.

Tribulus—A genus of branched annual or perennial herbs with some 25 species scattered in the tropics, subtropics, and temperate regions of the world. The name is from the Latinized Greek word tribolos, a 4-pointed weapon, in reference to the spiny or winged hard fruit. In the Hawaiian Islands represented by 1 indigenous species and 1 naturalized weedy species.

Tribulus cistoides L.

caltrop, nohu, nohunohu INDIGENOUS | SECURE

4.568 

Viola wailenalenae

Terrestial or epiphytic, erect, unbranched shrubs 10–50 cm tall. Leaves clustered toward the ends of the branches, slightly fleshy, elliptic ovate, 8–18 cm in length, glabrous. Flowers apparently all chasmogamous, petals white. Occurs in wet forest around the Alaka‘i Swamp on Kaua‘i at elevations of 1,220–1,410 m.

Viscaceae (MISTLETOE FAMILY)—The

Viscaceae, including familiar mistletoes and the Hawaiian species of Korthalsella, have now been merged into a broad concept of the Santalaceae.

Zygophyllaceae

4.569 

Tribulus cistoides

Prostrate to ascending perennial herbs with stems to several meters long. The fruit is spiny. Native to the Old World, now pantropical. In the Hawaiian Islands in coastal habitats at elevations from sea level to 50 m on all of the Northwestern Hawaiian Islands, except Gardner Pinnacles and Necker, and on all of the main islands.

(CREOSOTE BUSH FAMILY)

Herbs, shrubs, or rarely trees. Key features are the usually pinnate or 2-foliolate leaves and paired persistent stipules, flowers that have both a disc and distinct stamens with basal scales, and a single style. There are 26 genera and about 280 species that are predominantly tropical or subtropical in distribution, often in arid or saline places. Economically the family is of limited importance. Lignum vitae (Guaiacum) is the hardest and also the densest of woods. Creosote bush (Larrea) is a dominant shrub of both North and South American deserts and may be one of the oldest living plants in the world. A few species of Tribulus and

Monocotyledons Agavaceae (AGAVE FAMILY)—The genus Pleomele listed in the Agavaceae in the Manual is now included in the Asparagaceae. Araceae

(AROID FAMILY)

Herbs, often large and tuberous or rhizomatous and semi-epiphytic and epiphytic. Key features are an inflorescence as a fleshy spadix partially or completely enveloped by a spathe that is sometimes

petaloid or brightly colored. Foliage often with calcium oxalate crystals and milky sap. There are 105 genera and about 3,250 species with a pantropical distribution; the neotropical genera Philodendron and Anthurium comprise much of the diversity. Represented in the Hawaiian Islands by the arguably indigenous genus Spirodela. At times it has been suggested that Landoltia and Lemna might also be indigenous, but these are now considered to to be introduced. The Polynesians introduced ‘ape (Alocasia) and kalo or taro (Colocasia), and there are several naturalized ornamentals, including Zantedeschia, Epipremnum, and the aquatic herb water lettuce (Pistia).

Spirodela—Free-floating small plants on the water surface, with multiple roots and 5–11 nerves on each frond. A cosmopolitan genus of 5 species. The name comes from the Greek speira, cord or thread, and delos, visible, referring to the multiple roots on each frond. Represented in the Hawaiian Islands by the indigenous Spirodela polyrhiza. Occurs in taro paddies, slow-moving streams, ponds, and marshes in Kaua‘i, O‘ahu (Ko‘olau Mountains), Maui, and Hawai‘i. Possibly a naturalized introduction. Not pictured.

Arecaceae

(PALM FAMILY)

Trees, shrubs, and rarely lianas. Key features are the presence of fibrous roots, slender to massive solitary trunks or stems (rarely branched), and large pinnate or palmately divided leaves. There are 189 genera and about 2,400 species that are predominantly pantropical but with some representation in warmer temperate regions. Economically the family is important for food crops such as the coconut (Cocos) and dates (Phoenix), for wax (Copernicia), for palm oil (Elaeis), raffia (Raffia), rattan (Calamus), and betel (Areca). Other species provide thatch, vegetable ivory, wood, and other products. Many genera are widely cultivated, especially in tropical areas. In the Hawaiian Islands represented by the indigenous genus Pritchardia. The coconut palm (Cocus nucifera) is thought to be a Polynesian introduction. The king palm (Archontophoenix alexandrae) and the Chinese fan palm (Livistona chinensis) are widely naturalized in the Hawaiian Islands.

FLOWERING PLANTS

Pritchardia—A genus of medium-sized to

moderately tall solitary palms with fanshaped leaves found on tropical Pacific Islands. From 20 to 36 species are currently recognized by different authors, and the taxonomy is in need of revisionary work. Many species are known from very few individuals, and others survive only in cultivation. In the Hawaiian Islands, there are considered to be 22 endemic species, representing a single colonizing lineage. Many species are closely related, and the current treatment of these species is tentative. The name honors William T. Pritchard, nineteenth-century British consul to Fiji.

Pritchardia arecina Becc. loulu

ENDEMIC | SECURE

Pritchardia beccariana Rock loulu

ENDEMIC | SECURE

4.573 

4.571 

Pritchardia beccariana

Tall palms 16–19 m in height with slender trunks 20–30 cm in diameter and closely ringed. Leaf blades thin, coriaceous, lower leaf surface green, with scattered translucent scales, not waxy or glaucous, apices of leaf segment rigid, not drooping. Fruit 3.0–3.5 cm in length and 2.5–3.0 cm in diameter. An emergent wet forest species at elevations of 1,100–1,270 m on Hawai‘i (Mauna Loa).

Pritchardia hillebrandii

Medium-sized palms 6–7 m tall. Leaves rigid and thick, lower surface with scattered, pale-colored scales. Fruit globose and 2.0–2.2 cm in diameter. Occurs in mesic to wet areas at elevations of 30–600 m on windward Moloka‘i. Occasionally cultivated.

Pritchardia lowreyana Rock loulu

ENDEMIC | SECURE

Pritchardia hillebrandii (Kuntze) Becc. loulu, loulu lelo

ENDEMIC | SECURE

4.570 

Pritchardia arecina—habit

Medium-sized to tall palms, 10–12 m in height, trunks to 50 cm diameter. Lower surface of leaf blades densely covered with a pale yellow tomentum. Fruit large, globose-ovoid, 4.5 cm in length. Emergent in wet forests at 600–1,220 m elevation on the north slope of Haleakalā, East Maui.

4.574 

4.572 

Pritchardia hillebrandii—habit

Pritchardia lowreyana

Small palms 2–4 m tall. Leaf blades thick and coriaceous, 80–100 cm in length. Fruit large, ovoid-ellipsoid, 4–6 cm in length. Occurs in mesic valleys and inland cliffs at elevations of 130–920 m on Moloka‘i.

171

172

FLOWERING PLANTS

Pritchardia martii (Gaudich.) H. A. Wendl

Pritchardia minor Becc.

ENDEMIC | SECURE

ENDEMIC | SECURE

loulu, loulu hiwa

4.575 

loulu

Pritchardia martii—habit

Pritchardia minor

Medium-sized palms 6–12 m or more in height, with a slender trunk. Lower leaf surface covered with a soft yellowish gray or golden tomentum. Fruit small, 1–2 cm in length. Occurs in mesic to wet forest at elevations of 760–1,220 m Kaua‘i (western Nāpali Coast, Kōke‘e, and Waimea Canyon).

Pritchardia remota (Kuntze) Becc. loulu

ENDEMIC | ENDANGERED

4.578 

Pritchardia martii

Small to medium-sized palms 3–10 m tall. Lower surface of leaf blades silvery or with yellowish brown scales. Fruit large, ellipsoid, 4–5 cm in length. Scattered in wet forest on slopes, ridges, and cliffs or occasionally in mesic valleys at elevations of 360–610 m in the Ko‘olau Mountains, O‘ahu. The most common species of Pritchardia on O‘ahu.

(ASPARAGUS FAMILY)

Shrubs, rosette succulents, lianas, and herbs. There are two botanical views of the breadth of this family. In its narrow sense, it includes only a few genera and several hundred species, with Asparagus the most notable group. In the broader sense used here, it includes a range of what have been sometimes considered separate families such as the Agavaceae, Ruscaceae, Laxmanniaceae, and Hyacinthaceae. Under this broader concept, the family includes 103 genera and about 2,250 species and is cosmopolitan in distribution. In the Hawaiian Islands represented by the indigenous genus Pleomele.

4.577 

4.576 

Asparagaceae

Pleomele—A paleotropical genus of 40–50 species of shrubs or small trees. Some systematists place this group within Dracaena. The name is from the Greek pleon, many, and melon, apple, referring to the large number of fruit produced. Once placed in the Agavaceae and sometimes considered part of the Ruscaceae or in a smaller family concept of Dracaenaceae. In the Hawaiian Islands represented by 6 endemic species representing a single colonizing lineage. This genus was placed in the Agavaceae in the Manual.

Pleomele aurea (H. Mann) N. E. Brown hala pepe

ENDEMIC | SECURE

Pritchardia remota

Small palms 3–4 m tall, with slender, closely ringed trunks. Leaves dense and somewhat ruffled in appearance, lower surface with scattered, pale-colored scales. Fruit globose, 2 cm in length. Restricted to cliff bases and sea terraces at elevations of 200–800 m on Nīhoa (East and West Palm valleys) and Ni‘ihau (Ka‘ali Cliff). Known from 2 populations with about 680 individuals, but also in cultivation.

4.579 

Pleomele aurea—habit

FLOWERING PLANTS

Asteliaceae

(ASTELIA FAMILY)

Astelia menziesiana Sm.

pa‘iniu, kaluaha, pua‘akuhinia ENDEMIC | SECURE

Dwarf to large herbs, shrubs, and small trees, with some epiphytic. Key features are entire leaves, typically sessile, in a basal arrangement; flowers with 6 tepals. The family distribution is centered in New Zealand and Australia, with a single species in austral Chile, but absent from southern Africa. There are 4 genera and about 35 species.

4.580 

Pleomele aurea

Trees 6–12 m tall with only a few branches. Leaves semicoriaceous, 50–57 cm in length, 1.7–3.4 cm wide, tapering in their upper third. Inflorescence stout, 37–55 cm in length. Occurs in mesic forest and Pandanus forest at elevations of 140–1,070 m on Kaua‘i.

Pleomele hawaiiensis O. Deg. & I. Deg. hala pepe

ENDEMIC | ENDANGERED

Astelia—A genus of terrestrial or epiphytic perennial herbs, with 25 species found from Mauritius and Réunion to New Guinea, Australia, New Zealand and austral Chile and on many Pacific islands. The name is from the Greek a, without, and stele, column, referring to the acaulescent habit of the plants. In the Hawaiian Islands, there are 3 endemic species, representing a single colonizing lineage. Members of this genus are dioecious. This family was included in the Liliaceae in the Manual.

Astelia argyrocoma A. Heller ex Skottsb. pa‘iniu

ENDEMIC | SECURE

4.583 

Astelia menziesiana

Terrestrial or epiphytic perennial herbs with scaly leaves 1–5 cm wide. Flower color varies from green to yellow, brownish maroon, rose, or white. Male flowers in racemes 3–20 cm in length, female flower racemes 3–18 cm. In mesic to wet forest and bogs at elevations of 610–2,230 m on all of the main islands except Ni‘ihau and Kaho‘olawe. A variable plant with respect to the leaves and flower color, which in the past has led to the recognition of separate species. The plant pictured in habit is the form with the largest leaves from O‘ahu.

Cyperaceae 4.581 

(SEDGE FAMILY)

Pleomele hawaiiensis

Trees 5–6 m tall. Leaves semicoriaceous, 23–38 cm in length, 1.4–2.7 cm wide, gradually tapering in the upper third of the leaf. Inflorescences compact, 19–38 cm in length. Rare in dry or sometimes mesic forest at elevations of 300–860 m on the leeward side of Hawai‘i. Most recent estimates suggest a total population of 300–400 individuals. Threats have come most significantly from habitat loss, with feral grazing animals, alien plant species, and fire as secondary factors.

4.582 

Astelia argyrocoma

Epiphytic perennial herbs with narrow silvery leaves 0.4–1.0 cm wide. Flowers green to reddish brown, in racemes 6–10 cm long for males and 2–7 cm long for females. Occurs in wet forest and diverse mesic forest at elevations of 600–1,280 m on Kaua‘i.

Perennial grasslike or rushlike plants characteristically found in bogs, marshes, saturated soils, and riverine wetlands. Key vegetative features that separate this family from the grasses are solid and typically 3-angled stems, leaves arranged spirally in 3 ranks, and the sheathing leaf bases joined to the culm. Reproductive structure differs from the grasses in that each flower, often unisexual, is subtended by only a single bract. There are about 92 genera with some 4,450 species that are cosmopolitan in distribution.

173

174

FLOWERING PLANTS

Several genera have evolved C 4 metabolism. Economically the family is of little importance. Papyrus (Cyperus) was used by the Egyptians in papermaking. A few species have some ornamental value and are often used in artificial ponds. In the Hawaiian Islands represented by 15 indigenous genera: Bolboschoenus, Carex, Cladium, Cyperus, Eleocharis, Fimbristylis, Gahnia, Machaerina, Morelotia, Oreobolus, Rhyncospora, Schoenoplectiella, Schoenoplectus, Scleria, and Uncinia.

Bolboschoenus—A cosmopolitan genus

of 8 species of robust perennials found in aquatic habitats. This genus has been segregated from the larger genus Scirpus, where these species were formerly placed. The name is from the Greek bolbos, swelling, and schoinos, rush, referring to the tubers found at the base of the stems. In the Hawaiian Islands represented by 1 indigenous subspecies.

and tropical areas nearly worldwide. The subsp. paludosus is the form found in the Hawaiian Islands. A plant frequently encountered in mudflats and marshes, primarily in coastal sites on all of the main islands except Lāna‘i and Kaho‘olawe.

Carex meyenii Nees INDIGENOUS | SECURE

Carex—A large cosmopolitan genus of

herbaceous perennials comprising about 1,800 species distributed over a wide diversity of habitats but most common in wetlands. Plants monoecious, often with a woody rhizome. The name is from the Latin word for sedge. In the Hawaiian Islands represented by 4 endemic and 4 indigenous species representing 7 independent colonizing lineages.

Carex alligata Boott ENDEMIC | SECURE

4.586 

Carex meyenii

Densely tufted perennial, forming large clumps. Culms 3 angled, rigid, slender, 30–70 cm tall. Leaves numerous, stiff, narrowly linear, shorter than or as long as the culms. Stigmas 2, fruit flat. Occurs in dry to mesic forest at elevations of 180– 1,890 m on all of the main islands, but it is now extinct on Kaho‘olawe.

Carex montis-eeka Hillebr.

Bolboschoenus maritimus (L.) Palla

ENDEMIC | SECURE

kaluhā, bulrush

INDIGENOUS | SECURE

4.585 

4.584 

Bolboschoenus maritimus

Robust perennial 30–150 cm tall with creeping rhizomes that form tubers. A widely distributed plant in temperate

Carex alligata

Densely tufted perennials with short, woody rhizomes. Culms 3 angled, 60–150 cm tall. Leaves linear, nearly as long as the culms. Stigmas 2 and fruit flat, dull. Related to species from western North America. Occurs in wet forest mostly along stream banks or other wet places at elevations of 730–2,225 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i.

4.587 

Carex montis-eeka

Loose to moderately densely tufted perennials. Culms stiflly erect, 3 angled, 10–70 cm tall. Stigmas 3 and fruit with 3 angles. The linear leaves are unusually stiff. A bog plant found at elevations of 640–2,255 m on Kaua‘i, Moloka‘i, and Maui.

FLOWERING PLANTS

Carex wahuensis C. A. Mey. ENDEMIC | SECURE

Perennials 1–3 m tall with thick horizontal rhizomes, culm solitary, terete. Leaves cauline, coriaceous, linear, 60–90 cm in length. Widely distributed in the New World, on many Pacific islands, and also in Asia. Occurs in marshes, seeps, along streams, and around brackish ponds at elevations from sea level to 90 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i.

Cyperus—A cosmopolitan genus of small

4.588 

Carex wahuensis subsp. rubiginosa

Rhizomatous perennials forming large clumps. Culms 3 angled, 40–100 cm tall. Leaves coriaceous, up to 8 per culm. Stigmas 3, fruit 3 angled, constricted at angles. A common species occurring in dry to mesic forest, mesic and alpine shrubland, and sometimes mesic coastal forest at elevations of 250–2,550 m on all of the main islands except Ni‘ihau and Kaho‘olawe. There are 3 subspecies, with subsp. rubiginosa pictured.

to medium-sized annuals or herbaceous perennials with 3-ranked leaves. There are about 600 species found all over the world. The name is from the Greek kyperos, which means sedge. Many but not all species in the genus exhibit C 4 metabolism. Segregate genera once described for the Hawaiian Islands (i.e., Mariscus, Pycreus, Kyllinga, Torulinium) have now been returned to Cyperus. In the Hawaiian Islands represented by 9 endemic and 5 indigenous species, which come from 8 separate colonizing lineages. At least 11 other species have naturalized.

Cyperus hillebrandii Boeck ENDEMIC | SECURE

Cladium—A genus of 1–2 species of robust herbaceous perennials distributed in tropical and warm temperate regions of Eurasia, the Americas, and some Pacific islands. The plants are found in wetland habitats. The name is from the Greek kladion, small branch, referring to the shape of the inflorescences. In the Hawaiian Islands represented by 1 indigenous species.

compound, 2–5 cm in length. Occurs on ridges and gulches in dry to mesic forest at elevations of 250–1,980 m on Kaua‘i, O‘ahu (Wai‘anae Mountains), Lāna‘i, Maui, and Hawai‘i. Three subspecies are named, with subsp. hillebrandii pictured. Listed in the Manual as Mariscus hillebrandii.

Cyperus hypochlorus Hillebr. ‘ahu‘awa

ENDEMIC | SECURE

4.591 

Cyperus hypochlorus var. hypochlorus

Perennial sedge with short woody rhizomes clothed with reddish brown scales. Culms 50–130 cm tall, 0.5–1.6 cm in diameter at the base, 3 angled. Leaves several, coriaceous, linear, 3–15 mm wide, as long or longer than the culms. Occurs in mesic to wet forest at elevations of 350–1,280 m on all of the main islands except Ni‘ihau, Lāna‘i, and Kaho‘olawe. There are 2 named subspecies, with subsp. hypochlorus shown. Listed in the Manual as Mariscus hypochlorus.

Cyperus javanicus Houtt

‘ahu ‘awa, umbrella sedge INDIGENOUS | SECURE

Cladium jamaicense Crantz ‘uki, saw-grass

INDIGENOUS | SECURE

4.590 

4.589 

Cladium jamaicense

Cyperus hillebrandii var. hillebrandii

Perennials with short and typically thickened rhizomes, clothed with reddish brown basal sheaths. Culms 3 angled, 15–60 cm or more tall. Leaves stiff, linear, may be shorter or longer than the culms. Inflorescence umbelliform, simple or

4.592 

Cyperus javanicus

175

176

FLOWERING PLANTS

Coarse perennials with short rhizomes. Culms robust, 40–110 cm tall, obtusely 3 angled. Leaves numerous, usually longer than the culms. Inflorescence umbelliform, typically compound, 10–15 cm in length. Native to tropical Africa and Asia. Occurs in marshes, stream banks and ditches, and rocky coastal sites at elevations from sea level to 180 m on all of the main islands except Kaho‘olawe, as well as on Midway Atoll. The pounded stems of this plant were used by the Native Hawaiians to strain kava (‘awa). Listed in the Manual as Mariscus javanicus.

Cyperus odoratus L.

Cyperus laevigatus L.

Annuals or sometimes short-lived perennials. Culms erect, solitary or few together, 60–140 cm tall, 3 angled. Leaves several, linear, usually shorter than the culm, sheaths reddish brown. Inflorescence large and open, umbelliform,10–40 cm in length, 15–60 cm in width. Native to the West Indies, South America, Micronesia, Polynesia, and tropical Asia. Hawaiian plants are sometimes placed in the endemic subsp. auriculatum. Rare in low-elevation wet habitats such as margins of ponds, vernal pools, and taro paddies on O‘ahu; now considered to be extinct on Kaua‘i, Moloka‘i, Maui, and Hawai‘i. Listed in the Manual as Torulinium odoratum.

ehu‘awa, makaloa, umbrella sedge INDIGENOUS | SECURE

kili‘o‘opu, kiolohia, pu‘uka‘a, umbrella sedge INDIGENOUS | RARE

Cyperus polystachyos Rottb. INDIGENOUS | SECURE

4.594 

Cyperus odoratus

Cyperus pennatiformis Kük. ENDEMIC | ENDANGERED/EXTINCT?

4.593 

Cyperus laevigatus

Perennials 10–45 cm tall with creeping rhizomes clothed with brown scales. Culms arranged in a single row along the rhizome, 10–45 cm tall, 3 angled. Leaves reduced to bladeless sheaths, sheaths 2–3, brown to dark brown. Inflorescence with lateral axes longer than the primary axis, capitate with 1–12 sessile spikelets. Widespread in warm temperate and subtropical regions. Occurs on mud flats, edges of fresh, brackish, and saltwater ponds near the coast on Laysan, Ni‘ihau, O‘ahu, Moloka‘i, Maui, and Hawai‘i. On Ni‘ihau the leaves are woven into very fine, soft mats. Also used medicinally and for making hats.

the plants now appear to be reproducing successfully. The var. pennatiformis occurred from sea level to elevations of 1,200 m in coastal dunes, grasslands, and rarely mesic forest on Kaua‘i, O‘ahu, Maui, and Hawai‘i but is now possibly extinct. Listed in the Manual as Mariscus pennatiformis.

4.595 

Cyperus pennatiformis subsp. bryanii

Robust perennial sedge with culms 40–120 cm tall, sharply 3 angled. Leaves 3–5 per culm and somewhat coriaceous. Inflorescence umbelliform, moderately dense, 4–15 cm in length, 8–15 flowered. Rare in open sites of coastal dunes and today known only from Laysan. There are 2 named varieties, with the var. bryanii from Laysan shown. In the most recent survey, there were 5 populations and about 244 mature individuals of this variety on Laysan. Once decimated on the island by introduced rabbits,

4.596 

Cyperus polystachyos

Slender annuals or sometimes short-lived perennials with fibrous roots. Culms erect, 20–50 cm tall, 3 angled. Leaves few, basal, blades weakly plicate. Inflorescence open, forming an umbelliform corymb, sometimes contracted into a headlike cluster, 3–5 spreading bracts up to 30 cm in length, 8–40 flowered. A weedy plant that is found in tropical and subtropical areas worldwide. Common in a range of habitats from disturbed places to roadsides and in mesic and wet forests at elevations from sea level to 1,420 m on all of the main islands except Kaho‘olawe, as well as on Midway Atoll. Listed in the Manual as Pycreus polystachyos.

Eleocharis—A genus of slender annuals or herbaceous perennials found in wet habitats. Flowers perfect. About 200 species are recognized worldwide, with most distributed in temperate North and South America. The name is from the Greek helos, marsh, and charis, favor, referring to their preference for wetland habitats. Many but not all species in the genus exhibit C 4 metabolism. In the Hawaiian Islands, there are 2 indigenous species representing independent colonizing lineages and 2 naturalized species.

FLOWERING PLANTS

Eleocharis obtusa (Willd.) Schult. kohekohe, pīpīwai

INDIGENOUS | SECURE

Perennials with short rhizomes. Culms densely tufted, 5–60 cm tall. Leaves numerous, blades somewhat coriaceous, much shorter than culms. Sheaths persistent, thickly clothing the base of the culms. Inflorescence simple or compound, up to 4 cm in length and 7 cm in width, spikelets solitary to clustered. Widely distributed across the Pacific Basin in coastal areas and in shallow sandy soil or lava fields near the coast. In the Hawaiian Islands on Kure and Midway atolls, Laysan, French Frigate Shoals, and on all of the main islands except Kaho‘olawe. There are two subspecies, with subsp. umbellato-capitata pictured, with its inflorescence compacted into a single terminal head.

Gahnia—A genus of about 40 species of 4.597 

Eleocharis obtusa

Annual. Culms densely tufted, slender, 3–40 cm tall. Sheaths 1–6 cm in length. Spikelets densely many flowered. Widely distributed on the west coast of the United States and Canada. In the Hawaiian Islands occurring in wet areas, stream sides, and bogs at 390–1,900 m elevation on all of the main islands except Ni‘ihau, Kaho‘olawe, and Lāna‘i.

Fimbristylis—A cosmopolitan genus of

herbaceous perennials, often forming large rhizomatous clumps. Distributed in Australia, New Zealand, Malesia, Indochina, and the Pacific Islands. The name honors Henrik Gahn, an eighteenth-century student of Linnaeus. In the Hawaiian Islands, there are 2 endemic and 3 indigenous species, some of which have a preference for dry, open habitats. These taxa represent 3 independent colonizing lineages.

primarily basal, blades coriaceous, much longer than the culms, lower surface white powdery, involute. Inflorescence a moderately dense compound panicle, linear in shape, and 25–50 cm in length. Occurs in open sites and on ridges in mostly mesic forest at elevations of 300– 1,370 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Machaerina—A genus of rhizomatous

perennials with 2-ranked leaves and a preference for wetland sites. About 45 species are recognized around the world but with a concentration of diversity in Australia and New Zealand. The name is from the Latin machaera, sword, referring to the shape of the leaves. In the Hawaiian Islands represented by 2 indigenous species, with each from a separate colonizing lineage.

Machaerina angustifolia (Gaudich.) T. Koyama ‘uki

INDIGENOUS | SECURE

Gahnia beecheyi H. Mann ENDEMIC | SECURE

about 300 species of small to medium-sized annuals or herbaceous perennials. The name is from the Latin fimbria, fringe, and the Greek stylos, column, referring to the fringed nature of the style in some species. All species in the genus exhibit C 4 metabolism. The Hawaiian Islands have 1 endemic and 2 indigenous species, representing 3 independent colonizing lineages.

Fimbristylis cymosa R. Br. mau‘u ‘aki‘aki

INDIGENOUS | SECURE

4.600 

4.599 

4.598 

Fimbristylis cymosa subsp. umbellato-capitata

Gahnia beecheyi

Perennials with obliquely ascending rhizomes, thickly clothed with graybrown remnants of the leaf sheaths. Culms 60–120 cm tall. Leaves numerous,

Machaerina angustifolia

Perennials with short rhizomes. Culms smooth, 50–130 cm tall. Leaves pale green, overlapping in 2 ranks, primarily basal, as long as or slightly longer than the culms. Inflorescence a large compound panicle, 30–50 cm in length. Known from New Guinea, the Society Islands, and the

177

178

FLOWERING PLANTS

Hawaiian Islands. Occurs in wet forest and margins of bogs at elevations of 420– 2,070 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Morelotia gahniiformis Gaudich. ENDEMIC | SECURE

Machaerina mariscoides (Gaudich.) J. Kern ‘ahaniu, ‘uki

INDIGENOUS | SECURE

Densely tufted perennials usually forming large tussocks. Rhizomes ascending and branched, clothed with gray-brown remnants of old leaf sheaths. Culms rigid, 3–5 cm in length. Leaves numerous, densely tufted at apices of the rhizomes, irregularly 2 ranked, blades coriaceous. Inflorescence racemose, with 3–5 spikelets at 1–2 nodes. Occurs in bogs or open, cloudswept summits in wet forest at elevations of 640–2,320 m on Kaua‘i, O‘ahu, Moloka‘i, and Maui. Mosses, Viola, and stunted Vaccinium and Metrosideros often grow in the tussocks formed by this plant.

Rhynchospora—A genus of annuals or her4.602 

4.601 

Machaerina mariscoides

Perennials with short rhizomes clothed in reddish brown scales. Culms 45–90 cm tall, 3 angled. Leaves grayish green, coriaceous, largely basal, 2 ranked, upper ones longer than the culms. Inflorescence a compound panicle 15–30 cm in length. Distinguished from M. angustifolia by a shorter panicle with a zig-zag form. The species is native to New Guinea and several Pacific Island groups, with the Hawaiian plants placed into the endemic subsp. meyenii. Occurs in mesic to wet forest at elevations of 420–2,070 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Morelotia gahniiformis

Perennials with short, decumbent rhizomes clothed with brownish old leaf sheaths. Culms densely tufted, 20–70 cm tall, 3 angled. Leaves numerous, primarily basal, longer than the culms. Inflorescence an erect compound panicle, 10–35 cm in length. New Zealand plants, once considered to be identical, are now split off as a separate species. Occurs in dry forest, mesic forest, subalpine shrubland, and lava fields at elevations of 520–2,380 m on Kaua‘i (Wahiawa), Moloka‘i, Lāna‘i, Maui, and Hawai‘i. Listed in the Manual as Gahnia gahniiformis.

Rhynchospora rugosa (Vahl) Gale pu‘uko‘a, kuolohia, beak-rush INDIGENOUS | SECURE

Oreobolus—A genus of tussock-forming

herbaceous perennials in boggy habitats. The genus includes 14 species distributed in the high mountains of Central and South America, Malesia, Australia, and the Hawaiian Islands, as well as subantarctic regions of South America. The name is from the Greek oreos, mountain, bolos, bump, referring to the habitat and shape of the plants. In the Hawaiian Islands, there is 1 endemic species.

Oreobolus furcatus H. Mann ENDEMIC | SECURE

Morelotia—A small genus with 2 species of rhizomatous herbaceous perennials forming large tussocks. Native to New Zealand and the Hawaiian Islands. The origin of the name is uncertain. These species were formerly placed in the genus Gahnia. Represented in the Hawaiian Islands by 1 endemic species.

baceous perennials with usually tufted stems, widely distributed in the tropics and subtropics of the world. About 250 species are recognized. The name is from the Greek rhyncos, beak, and spora, seed, referring to the shape of the fruit. In the Hawaiian Islands, there are 3 indigenous and 2 naturalized species, representing 3 independent colonizing lineages.

4.603 

Oreobolus furcatus

4.604 

Rhynchospora rugosa var. lavarum.

Perennials without conspicuous rhizomes. Culms tufted, 20–70 cm tall. Leaves few, linear, basal, 8–30 cm in length, 0.8–1.5 mm in width. Inflorescence of 2–5 small panicles with the lateral branches exceeding the main axis. Native to the neotropics and reaching the Hawaiian Islands, where it occurs in wet forests, bogs, and occasionally mesic forests at elevations of 280–1,620 m on

FLOWERING PLANTS

all of the main islands except Ni‘ihau, Kaho‘olawe, and Lāna‘i. Hawaiian plants are in the indigenous subsp. lavarum.

Rhynchospora sclerioides Hook. & Arn. kuolohia, beak-rush

INDIGENOUS | SECURE

77 species are found all over the world, including the familiar tule or bulrush in the United States. These species were formerly considered in a broader concept of the genus Scirpus. The name is from the Greek schoinos, rush, and plektos, twisted, referring to the shape of the disintegrated persistent fibers. The genus is often included in Scirpus. In the Hawaiian Islands represented by the indigenous Schoenoplectus tabernaemontani and 1 naturalized species. Listed in the Manual as S. lacustris.

hooked barbs. Plants monoecious. About 54 species are known from South America, New Zealand, Australia, western Malesia, and the Hawaiian Islands. The name is from the Latin uncus, barb, in reference to the hooks on the inflorescence. In the Hawaiian Islands, there are 2 indigenous species representing separate colonizing lineages.

Uncinia uncinata (L. fil.) Kükenth. birdcatching sedge

INDIGENOUS | SECURE

Scleria—A pantropical genus of mostly

small to large perennial sedges or occasionally annuals. Plants monoecious. About 250 species are known. The name is from the Greek skleros, hard, referring to the texture of the small fruit. In the Hawaiian Islands represented by 1 indigenous species.

Scleria testacea Nees nutrush

INDIGENOUS | SECURE

4.605 

Rhynchospora sclerioides

Rhizomatous perennials with a short rhizome covered with brown fibers. Culms stout, 40–90 cm tall, 3 angled. Leaves numerous, primarily basal, as long or longer than the culms, 17–18 mm in width. Inflorescence of 5–9 small panicles, overall elongate and narrow, 15–60 cm in length. Native to Polynesia. Occurs in mesic forest, on ridges, and on grasslands at elevations 90–1,220 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Schoenoplectiella—A genus of about 26 species of annuals or herbaceous perennials that range from small sedges to tall bulrushes. Often treated within Schoenoplectus or more broadly in the past in Scirpus. The name is from the Greek schoinos, rush; plektos, twisted, referring to the shape of the disintegrated persistent fibers; and ella, the diminutive, in reference to Schoenoplectus. Represented in the Hawaiian Islands by Schoenoplectiella juncoides. Listed in the Manual as Schoenoplectus juncoides. Schoenoplectus—A cosmopolitan genus of annuals or herbaceous perennials with the leaves reduced to basal sheaths. About

4.607 

4.606 

Scleria testacea

Perennials with horizontal woody rhizomes. Culms densely tufted along the rhizomes, 60–90 cm tall, 3 angled. Leaves linear, 30–50 cm in length. Inflorescence a narrow compound panicle. The dingy white fruit of this plant is characteristic. Widely distributed in the neotropics. Known only from bogs on West Maui (Pu‘ukukui) and low-elevation wet sites in the area around Hilo.

Uncinia—A genus of herbaceous perenni-

als with the inflorescence axis possessing

Uncinia uncinata

Perennials with short rhizomes. Culms densely tufted, 20–60 cm tall, 3 angled. Leaves linear, typically longer than the culms. The fruit with attached covering and stalk is easily detached and fall as a unit. Native to New Zealand and the Hawaiian Islands. Occurs in wet forest and sometimes in bogs at elevations of 1,040–2,100 m on Kaua‘i, Moloka‘i, Maui, and Hawai‘i. The fruit is easily transported by feathers or skin of birds.

Hemerocallidaceae—The Hawaiian genus Dianella, formerly placed in the Hemerocallidaceae, is now included in a broader concept of the Xanthorrhoeaceae. Hydrocharitaceae (FROG’S BIT FAMILY)

Aquatic herbs, partly or totally submerged. Key features are the typical presence of unisexual flowers, an inferior

179

180

FLOWERING PLANTS

ovary, the spathe or paired floral bracts, and the male flowers often breaking free from submerged inflorescences and floating to the surface. There are 168 genera and about 120 species that occur in temperate and tropical regions. Economically the family is of no importance. Some species of Anacharis and Vallisneria are used as aquatics in fish tanks. Represented in the Hawaiian Islands by the indigenous saltwater aquatic Halophila.

Halophila—A genus of submerged marine

aquatics with creeping stems that root at the nodes. There are 10 species widely distributed, except not found in Europe and North and South America, although present in the Caribbean. The name is from the Greek holos, sea, and philos, fond of, referring to the marine habit of the plants. Like other seagrasses, Halo­ phila meadows support a rich community of associated organisms on the leaf blades and in associated sediments, as well as providing a food source for the Hawaiian green turtle. In the Hawaiian Islands represented by 1 endemic species and 1 indigenous species.

Halophila hawaiiana Doty & B. C. Stone seagrass

ENDEMIC | SECURE

Iridaceae

(IRIS FAMILY)

Perennial herbs or rarely subshrubs. Key features are the presence of rhizomes, bulbs, or corms, leaves that are usually flattened in a single plane, showy flowers with a perianth of 6 petaloid sepals in 2 whorls, and a capsular fruit often bearing large seeds. There are 70 genera and about 2,000 species that are cosmopolitan but with a high concentration of diversity in southern Africa. Economically the genus is important for the large number of ornamentals, such as Iris, Crocus, Freesia, Ixia, Tigridia, and Gladiolus. In the Hawaiian Islands represented by the indigenous genus Sisyrinchium. Species of Aristea, Crocosmia, Watsonia, and Trimezia have naturalized.

Sisyrinchium—A genus of perennial herbs, occasionally annuals, arising from rhizomes or from a crown with fleshy fibrous roots. There are about 140 species found in North and South America and the Hawaiian Islands. The name is from the Greek sisyrinchion, a common name for the barbary nut (Iris sisyrinchium). In the Hawaiian Islands represented by 1 endemic and 2 introduced species. Several species of this genus have horticultural merit and are often cultivated.

Sisyrinchium acre H. Mann

mau‘u hō‘ula ‘ili, mau‘u lā‘ili, blue-eyed grass

stems broadly winged, lacking cauline leaves. Flowers several per flowering stem, yellow. Occurs in open, dry to boggy places and in subalpine shrubland at elevations of 1,550–2,950 m on East Maui and Hawai‘i. The juice and leaves yield a blue dye that the Native Hawaiians used in tattooing.

Joinvilleaceae

(JOINVILLEA FAMILY)

Coarse, grasslike rhizomatous herbs. Key features are the linear leaves with many parallel veins well distributed along the stem. Small flowers produced on a terminal, many-branched inflorescence, and with the fruit a small, 3-angled drupe. The sole genus is found in western Malesia eastward to the Hawaiian Islands and Samoa. Economically the family is of no importance. The genus was once included as a second member of the Flagellariaceae but differs from Flagellaria in lacking tendrils.

Joinvillea—A genus of 2 species of erect

perennial herbs occurring on Pacific islands. The name honors Prince de Joinville, the third son of King Louis Phillipe of France. In the Hawaiian Islands represented by a rare endemic subspecies of 1 taxon.

Joinvillea ascendens Gaudich. ex Brongn & Gris ‘ohe

INDIGENOUS | RARE

ENDEMIC | RARE

4.608 

Halophila hawaiiana

Submerged saltwater perennials with unisexual flowers. Pairs of leaves occur on petioles along a rhizome rooted in sand. Leaves 1.8–5.0 mm wide, obovate to spatulate. Male and female flowers are produced infrequently on separate plants. Branching leads to intertwined plants in a turf with runners colonizing adjacent substrate. Found subtidally at 0.5–4.0 m depth in muddy or sandy areas of fishponds and sandy reef flats on Midway and Pearl and Hermes atolls, Kaua‘i, O‘ahu, Moloka‘i, and Maui.

4.609 

Sisyrinchium acre

Perennial herbs 15–30 cm tall with fibrous roots. Leaves narrow, basal, about half as long as the flowering stems. Flowering

4.610 

Joinvillea ascendens subsp. ascendens

Erect grasslike herbs 1.5–5.0 m tall. Leaf blades narrowly elliptic, 45–80 cm in length, 4.5–16.0 cm in width, plicate with

FLOWERING PLANTS

7–13 pleats, both surfaces with scattered bristles. Flowers in a large, branched inflorescence. The small fruit is bright orange. Native to New Caledonia, the Caroline Islands, and Western Samoa. Rare in wet forest and along intermittent streams at elevations of 300–1,250 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i. The Hawaiian taxon represents the endemic subsp. ascendens.

Juncaceae

(RUSH FAMILY)

Perennial or sometimes annual herbs. Key features are the presence of a rhizomatous rootstock, mostly basal, tufted, grasslike leaves, a perianth of papery segments usually in 2 series of 3, and the fruit a capsule. There are 7 genera and about 430 species mostly confined to temperate regions of the world and rare in the tropics. Like the Cyperaceae, these taxa are typically found in bogs, marshes, saturated soils, and riverine wetlands. Economically the family is of little importance. A few species have some ornamental value, especially in aquatic habitats. Some are used in weaving of hats, chair seats, and other products. In the Hawaiian Islands represented by the indigenous genus Luzula and 6 introduced species of Juncus.

Luzula—A genus of usually tufted peren-

nial herbs with soft leaves, often hairy on the margins. There are 108 species worldwide but confined mostly to north temperate areas. The name may have come from the Italian lucciole, glow worm, perhaps because of the appearance of the hairs when dew covered. In the Hawaiian Islands represented by 1 endemic species.

Luzula hawaiiensis Buchenau wood rush

ENDEMIC | SECURE

4.611 

Luzula hawaiiensis var. oahuensis

Low, tufted perennial herbs with fibrous roots, stems erect. Leaves basal, linear, 5–30 cm in length, margins densely pubescent. Inflorescence cymose, terminal, with 1–4 leaflike bracts 2–10 cm in length. Occurs in open or rocky places, slopes, and ridges in mesic forest and shrubland as well as in subalpine forest, shrubland, and grassland at elevations of 730–2,560 m on all of the main islands except Ni‘ihau and Kaho‘olawe. There are 3 varieties that differ in habit, pubescence, and inflorescence type. Pictured is the var. oahuensis with a range restricted to the Wai‘anae Mountains, O‘ahu.

Lemnaceae

Anoectochilus—A genus of about 30 species of terrestrial creeping or decumbent orchids with small chartreuse or white flowers. The name is from the Greek anoiktos, open, and cheilos, lip, referring to the shape of the lip of the flower. In the Hawaiian Islands represented by 1 endemic species confined to wet, shady places.

Anoectochilus sandvicensis Lindl. jewel orchid

ENDEMIC | RARE

(DUCKWEED FAMILY)—

The genera Lemna and Spirodela listed in the Manual in the Lemnaceae are now included in the Araceae.

Liliaceae (LILY FAMILY)—Two Hawaiian genera formerly placed in the Liliaceae (sensu lato), Astelia and Dianella, have now been placed in the Asteliaceae and Xanthorrhoeaceae, respectively. Orchidaceae

(ORCHID FAMILY)

Perennial herbs, both epiphytic and terrestrial, occasionally saprophytic. Key features are the highly complex anatomy of the flowers, the often highly specialized nature of pollination, the capsular fruit with numerous, minute seeds, and the mycorrhizal association necessary for germination. The largest family of monocots and second largest of flowering plants with 779 genera and about 22,500 species; the greatest diversity of species occurs in the tropics. Economically the family is important for the large number of genera in cultivation and for the production of vanilla from the dried fruit of Vanilla. In the Hawaiian Islands represented by only 3 indigenous genera with 1 species each. Perhaps the lack of suitable pollinators or absence of appropriate mycorrhizal symbionts may explain the paucity of native orchids. Arundina, Epidendrum, Phaius, Polystachya, Spathoglottis, and Zeuxine have species that have naturalized, with some of these becoming weedy. Orchids are widely cultivated in the Islands.

4.612 

Anectochilus sandvicensis

Decumbent perennial herbs up to 50 cm tall arising from rhizomes, adventitiously rooting at the nodes. Leaves dark green, glossy, 4–8 cm in length, restricted to the upper part of the stems. Flowers few to numerous in loose spikes, 5–15 cm in length. Rare in wet shady places, usually terrestrial but occasionally epiphytic on tree ferns, in wet forest at elevations of 275–1,710 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Liparis—A cosmospolitan genus of about 250 species of small to medium-sized perennial orchids that may be either terrestrial or epiphytic. The name is from the Greek liparos, smooth, referring to the shiny surface of the leaves. In the Hawaiian Islands represented by 1 endemic species.

181

182

FLOWERING PLANTS

Liparis hawaiensis H. Mann

‘awapuhi-a-Kanaloa, twayblade ENDEMIC | RARE

Platanthera—A genus of about 100 species of terrestrial orchids distributed mostly in north temperate regions of the Northern Hemisphere but also in Malesia and on some Pacific islands. Formerly included in Habenaria or Orchis, 2 closely related genera, but distinguished from these by the distinctive stigma lobes. The name is from the Greek platys, broad, and anthera, anther, referring to the gap between the anther sacs. Represented in the Hawaiian Islands by 1 endemic species.

Platanthera holochila (Hillebr.) Kraenzl. fringed orchid

ENDEMIC | ENDANGERED

4.613 

with alternate leaves that appear to be spirally arranged. Plants dioecious. The large fleshy fruit are berries or multichambered drupes and often resemble small pineapples. There are 4 genera and about 825 species found primarily in the paleotropics, tropics, and reaching New Zealand. Economically the family is important for the leaves of various species of Pandanus, which are used for thatch, matting, and clothing. One species with edible leaves is widely used in flavoring soups and candies, particularly in Southeast Asia. Multiple cultivars have been selected for edible fruits throughout Micronesia and form an important staple of local diets in many areas of the Pacific. Species of both Pandanus and Freycinetia are cultivated as ornamentals in tropical areas. Represented in the Hawaiian Islands by the indigenous genera Freycinetia and Pandanus.

Freycinetia—A genus of woody climbers, often with long, adventitious roots. About 300 species are recognized in Southeast Asia, Malesia, Australia, Norfolk Island, New Zealand, and many Pacific islands. The name honors Louis de Freycinet, an eighteenth-century ship captain and explorer. In the Hawaiian Islands represented by 1 indigenous species.

Liparis hawaiensis—habit

Freycinetia arborea Gaudich. ‘ie, ‘ie‘ie

INDIGENOUS | SECURE

4.615 

4.614 

Liparis hawaiensis

Erect terrestrial or epiphytic perennial herbs, 5–15 (30) cm tall. Leaves 1–2 arising from each pseudobulb, crowded along the rhizome, ovate, 1–8 cm in length. Flowers pale green, in erect terminal racemose inflorescences 15–30 cm in length. Rare on moss-covered trees, seasonally wet bare ground, and bogs in mesic to wet forest at elevations of 490–1,530 m on all of the main islands except Ni‘ihau and Kaho‘olawe. The most commonly encountered native orchid.

Platanthera holochila

Terrestrial deciduous herbs up to 60 cm tall arising from subterranean tubers. Leaves pale green, membranous, lanceolate to ovate, 4–12 cm in length. Flowers greenish yellow, in open spikes subtended by a series of linear-lanceolate leaflike bracts. Rare in bog hummocks or wet forests surrounding bogs or in mesic forest and subalpine shrubland at elevations of 600–1,830 m on Kaua‘i, O‘ahu (extinct), Moloka‘i, and Maui. Critically endangered and known from only 3 populations totaling 26 individuals. Threats come from cattle, feral pigs, and alien plant species.

Pandanaceae

(SCREW PINE FAMILY)

Trees, shrubs, or lianas. Key features are the often woody, palmlike habit, aerial support roots, and often twisted stems

4.616 

Freycinetia arborea

FLOWERING PLANTS

Woody climbers in trees or sometimes sprawling on the ground. Stems with long aerial roots that help support the plants. Leaves coriaceous, linear-lanceolate, 40–80 cm in length, prickles on the margins. Inflorescences terminal with colored, leafy bracts. Fruiting spadices 8–10 cm in length, berries ripening red. Widespread in Polynesia and also in New Caledonia. Common in mesic to wet forest at elevations of 300–1,500 m on all of the main islands except Ni‘ihau and Kaho‘olawe. The bright-colored bracts subtending the inflorescence are often eaten by rats, which pose a major threat to their survival in some areas. Pollination is effected by birds. The 4 original pollinating bird species are all extinct or endangered, and the introduced Japanese white-eye (Zosterops japonicus) is now the principal pollinator. The Native Hawaiians utilized the stem fibers for cordage, while the roots were woven into baskets and fish traps and plaited into helmets worn by the chiefs in battle.

Pandanus—A genus of often palmlike trees and shrubs that are terrestrial, epiphytic, or semiaquatic with prop roots extending from the base of the stem. There are about 520 species of the paleotropics, with the greatest diversity in species occurring in Malesia and Madagascar. The name is from the Amboinese word pandan. In the Hawaiian Islands represented by 1 widespread indigenous species.

Pandanus tectorius Parkinson hala, pū hala, screw pine INDIGENOUS | SECURE

4.617 

Pandanus tectorius

Small trees to 10 m tall with mostly erect trunks and many branches supported by numerous thick-branched prop roots. Leaves crowded near the branch ends, coriaceous, 80–180 cm in length. Inflorescences terminal, monoecious, with slender, short-lived white male inflorescences and female inflorescences resembling a small pineapple. Native to Polynesia, Micronesia, Melanesia, New Guinea, New Caledonia, northern Australia, the Philippines, the Moluccas, and Java. Occurs in coastal sites or low-elevation slopes of mesic valleys at elevations from sea level to 600 m on all of the main islands except Kaho‘olawe. This tree had many uses for the Native Hawaiians. Woven leaves were used to make sails for the Polynesian voyaging canoes. The old dry fruit was used as brushes to dye kapa, and fresh ones were used in lei making. In the Hawaiian tradition, hala lei are used for funerals and for commemorating the dead, although modern use is more decorative than symbolic. Leaves are woven into mats, bags, and purses. Roots can be retted to obtain a cordage fiber. The seeds are edible, but the fruit must first be cooked to eliminate calcium oxalate crystals. Fossil material recently collected in the Hawaiian Islands suggests the indigenous nature of this plant, which has often been considered a Polynesian introduction.

plants, for food sources such as wheat (Triticum), rice (Oryza), sugarcane (Saccharum), oats (Avena), corn (Zea), rye (Secale), and sorghum (Sorghum). Many species are important for providing shelter in the form of thatch, framing, and matting; in industry for the production of ethyl alcohol; for turf (Poa, Festuca, and Agrostis); and for ornamentals such as bamboo. In the Hawaiian Islands represented by the monotypic endemic genus Dissochondrus and the indigenous genera Agrostis, Calamagrostis, Cenchrus, Deschampsia, Dichanthelium, Digitaria, Eragrostis, Festuca, Isachne, Ischaemum, Lepturus, Panicum, Poa, Sporobolus, and Trisetum. Chrysopogon, Heteropogon, Paspalum, and Schizostachyum are possibly indigenous or early introductions. At least 47 other genera have naturalized, and many of these are major invasives with significant ecological impacts (see chapter 7).

Poaceae

bentgrass

(GRASS FAMILY)

Annual or perennial herbs (some bamboos woody) that are morphologically complex and possess a terminology peculiar to this family. Key features are the unique inflorescence, called a spikelet, with its component parts (i.e., glume, lemma, and palea), and the flower itself with 2 lodicules (or reduced perianth parts), the stamens usually in a whorl of 3, and a pistil with a superior ovary and 2 brushy styles. Fruit a caryopsis (or in some bamboos, a nut or berry), and seeds with copious endosperm. Probably the fourth largest family of flowering plants, with about 715 genera and 10,550 species. Many genera exhibit C 4 metabolism, and both native and alien species with this ecophysiological trait are dominant at lower elevations in the Hawaiian Islands. Economically this is probably the most important of all the families of flowering

Agrostis—A genus of annual or herbaceous perennial grasses with slender culms and an open to contracted inflorescence. There are about 175 species found in warm to cool temperate areas around the world. The name is from the Greek agrostis, an old name for a weedy fodder plant. In the Hawaiian Islands represented by 1 endemic species.

Agrostis sandwicensis Hillebr. ENDEMIC | SECURE

4.618 

Agrostis sandwicensis

183

184

FLOWERING PLANTS

Erect, tufted perennial grasses, culms solitary or in small clusters, 15–30 cm tall. Blades linear, 5–10 cm in length, 1–2 mm in width, scabrous. Inflorescence paniculate, cylindrical, 8–12 cm in length, with a purple tinge, spikelets numerous. Occurs in subalpine cinder fields on Kaua‘i, Maui, and Hawai‘i and in open bogs at elevations of 1,190–3,540 m. Reported by Hillebrand from O‘ahu in 1888.

Calamagrostis—A genus of large herba-

ceous perennial grasses with narrow glabrous leaves and an open to contracted inflorescence. Plants may be caespitose, rhizomatous, or stoloniferous. There are about 250 species found mostly in temperate regions of the world but also with some representation in tropical montane areas. The name is from the Greek kalamos, cane, and the genus Agrostis because it is somewhat intermediate between 2 kinds of grasses. In the Hawaiian Islands, there are 2 endemic species representing independent colonizing lineages. Calamagrostis expansa is a rare endemic occurring on Maui and the Kohala Mountains, Hawai‘i.

Calamagrostis hillebrandii (Munro ex Hillebr.) Hitchc.

Agrostis sandwicensis but differing from the former in the presence of a single floret and a subapically attached awn, and from the latter in having long hairs on the axis of the spikelet. Occurs in open bogs at elevations of 1,220-2,990 m at Pu‘ukukui on West Maui and once reported by Hitchcock in 1922 from central Moloka‘i.

Cenchrus—A genus of annual or herbaceous perennial grasses traditionally including species with the involucres, which are formed of fused bristles or spines subtending and partially covering the spikelets, covered with spines or bristles; these fall as a unit. Recent molecular studies have now placed all species of Pennisetum within Cenchrus. There are 23 species found in the New World and across the Pacific Basin to tropical Asia and Australia. Species exhibit C 4 metabolism. The name is from the Greek kenchros, a small millet or cereal grass used by the ancient Greeks. In the Hawaiian Islands represented by 1 endemic and at least 10 naturalized species.

5–25 mm in width, upper surface scabrous, lower surface glabrous. Inflorescence composed of racemes 5–10 cm in length, spikelets subtended by a bristly bur. The var. agrimonioides occurs on dry, rocky slopes and ridges in partial shade at elevations from sea level to 760 m on O‘ahu, Lāna‘i, and Maui. The var. laysanensis once found on Kure and Midway atolls and Laysan is now extinct.

Chrysopogon—A

genus of perennial grasses with tufted culms. About 44 species are distributed in the paleotropics and warm temperate Asia and Australia. The name comes from the Greek chrysos, gold, and pogon, beard, referring to the golden hairs that are present at the base of the glume on the sessile spikelets. In the Hawaiian Islands represented by 1 indigenous species.

Chrysopogon aciculatus (Retz.) Trin.

mānienie ‘ula, pi‘ipi‘i, golden beardgrass INDIGENOUS? | SECURE

Cenchrus agrimonioides Trin. kāmanomano, sandbur

ENDEMIC | ENDANGERED/EXTINCT

ENDEMIC | VULNERABLE

4.621 

4.619 

Calamagrostis hillebrandii

Perennial grass with erect culms and short rhizomes, 30-50 cm tall. Appears similar to Deschampsia nubigena and

4.620 

Cenchrus agrimonioides var. agrimonioides

Perennial grasses with robust culms swollen at the nodes, 30–200 cm tall. Blades flat or folded, 20–40 cm in length,

Chrysopogon aciculatus

Perennial grasses with creeping stolons, covered with imbricate scalelike old sheaths. Blades 2–8 cm in length, 3–5 mm in width, glossy, margins scabrous. Inflorescence paniculate, narrowly elliptic, 3–6 cm in length, purple, branches whorled. Native to tropical Malesia, Australia, and arguably the Pacific Basin but possibly an early and inadvertent Polynesian introduction. Found on dry exposed slopes and ridges at elevations of 15–520 m on all of the main islands except Ni‘ihau, Kaho‘olawe, and East Maui.

FLOWERING PLANTS

Deschampsia—A genus of tufted annual or perennial grasses with inflorescences open or contracted and usually with slender wiry branches and pedicels. About 30 species are found in temperate and arctic areas worldwide and some representation in the montane tropics. The name honors Jean-Louis-Auguste Loiseleur-Deslongchamps, an eighteenth-century French naturalist and doctor. In the Hawaiian Islands represented by 1 endemic species.

Dichanthelium cynodon (Reichardt) C. A. Clark & Gould ENDEMIC | SECURE

Deschampsia nubigena Hillebr. hairgrass

ENDEMIC | SECURE

4.623 

Dichanthelium cynodon

Herbaceous perennial grasses forming dense mats or tussocks to 60 cm in diameter, culms erect, many branched, 8–20 cm tall. Blades flat, narrowly lanceolate, 1.5–3.5 cm in length, 1–3 mm in width. Inflorescence paniculate, 1.5–3.0 cm in length. Forming mats or tussocks in open bogs at elevations of 1,220–1,950 m on Kaua‘i, Moloka‘i, and Maui.

Digitaria—A cosmopolitan genus of about 250 species of annual or perennial grasses, usually stoloniferous. While many species are aggressive weeds, a few are important grain crops called fonio in Africa. Species exhibit C4 metabolism. The name comes from the Latin digitus, finger, referring to the palmately arranged branches of the inflorescence of most species. In the Hawaiian Islands represented by Digitaria setigera, which may represent an early colonization or an inadvertent introduction by the Polynesians. Dissochondrus—An endemic, monotypic Hawaiian genus of grasses with short creeping rhizomes and a narrow, cylindrical, spikelike inflorescence. The name is from the Greek dissos, double, and chondros, cartilage, referring to the 2 cartilaginous fertile florets. Species exhibit C 4 metabolism. Represented in the Hawaiian Islands by 1 endemic species. Closely related to Setaria.

Dissochondrus biflorus (Hillebr.) Kuntze ex Hack. ENDEMIC | RARE

Dichanthelium isachnoides (Munro ex Hillebr.) C. A. Clark & Gould ENDEMIC | SECURE

4.622 

Deschampsia nubigena

Tufted perennials, culms stiff, 30–100 cm tall, glabrous. Blades folded, linear, glabrous, margins scabrous, 5–30 cm in length, 0.5–1.5 mm in width. Inflorescence paniculate, open oblong-ovate, 5–30 cm in length. Occurs in mesic and wet forest, bogs, subalpine grassland, subalpine shrubland, and cinder fields at elevations of 600–2,830 m on Kaua‘i, Moloka‘i, Maui, and Hawai‘i. A variable species.

Dichanthelium—A genus, formerly included in Panicum, of tufted or tussock-forming herbaceous perennial grasses. About 72 species are found primarily in the New World. The name is from the Greek dicha, two or against, and anthele, an inflorescence type, perhaps referring to two different inflorescence types produced by many species in this genus. In the Hawaiian Islands, there are 4 endemic species, usually found at higher elevations, which represent a single colonizing lineage.

4.624 

Dichanthelium isachnoides

Herbaceous perennial grasses forming tussocks, culms stout, tufted, 15–40 cm tall. Leaves closely imbricate. Blades 0.8– 6.5 cm in length, 3.5–8.0 mm in width, ciliate with long, stiff hairs along the margins. Forms tussocks in open bogs at elevations of 910–1,850 m on Kaua‘i, Moloka‘i, and Maui.

4.625 

Dissochondrus biflorus

Perennial grasses with short creeping rhizomes, culms 60–120 cm tall, compressed. Blades flat, firm, 15–30 cm in length, 10–20 mm in width. Inflorescence paniculate, narrowly cylindrical and spikelike, 10–20 cm in length. Occurs on slopes in diverse mesic forest at elevations

185

186

FLOWERING PLANTS

of 490–1,050 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Now extinct on Kaua‘i.

Eragrostis—A genus of annuals or herbaceous perennials with open or contracted spikelike inflorescences. There are about 350 species worldwide in temperate and tropical areas. Many species are cosmopolitan weeds. The name is from the Greek eros, loving, and Agrostis. Species exhibit C 4 metabolism. Teff (Eragrostis tef ) is a crop of commercial importance in Ethiopia and Somalia, where it is used to make traditional breads. In the Hawaiian Islands, there are 8 endemic and 1 indigenous species representing 2 independent colonizing lineages; there are also 13 naturalized species.

Eragrostis atropiodes Hillebr. ENDEMIC | SECURE

Eragrostis grandis Hillebr. lovegrass

ENDEMIC | SECURE

4.627 

Eragrostis grandis

Perennial grasses with the culms forming large tufts, up to 200 cm tall. Blades flat, 5–10 mm in width. Inflorescence open and paniculate, up to 50 cm in length. Common on moist to wet slopes in ravines and mesic forest, usually in partial shade at elevations of 170–2,040 m on all of the main islands except Ni‘ihau.

Eragrostis variabilis (Gaudich.) Steud. kāwelu, kalamālō, ‘emoloa, lovegrass ENDEMIC | SECURE

forest, and exposed sites and ridges from sea level to 1,130 m on Kure, Midway, and Pearl and Hermes atolls, Lisianski, Laysan, Nīhoa, and all of the main islands. A variable species.

Festuca—A genus of caespitose herbaceous perennial grasses with an open or dense inflorescence. There are about 4,500 species worldwide. The name is from the Latin festuca, culm or stem. In the Hawaiian Islands represented by 3 endemic species and 2 naturalized species. Festuca hawaiiensis is a rare or possibly now extinct endemic known only from moderately high elevations at ‘Ulupalakua on Maui and Pu‘uhuluhulu and Hualālai on Hawai‘i. Festuca aloha is a vulnerable endemic from Kaua‘i, while Festuca molokaiensis is a local endemic from Kūpā‘ia Gulch, Moloka‘i. Heteropogon—A genus of 8 species of annuals or herbaceous perennial grasses with a solitary racemose inflorescence found in southern Europe, Africa, and the tropics. The name is from the Greek heteros, different, and pogon, beard, referring to the 2 kinds of spikelets found in the inflorescence. Species exhibit C 4 metabolism. In the Hawaiian Islands represented by 1 possibly indigenous species.

Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult.

pili, pili grass, tanglehead, twisted beardgrass INDIGENOUS/POLYNESIAN INTRODUCTION  |  SECURE

4.626 

Eragrostis atropioides

Perennial grasses with loose tufts, rhizomes hard and scaley and up to 7 cm in length. Culms stiff and erect, 100–150 cm tall. Leaves flat, 30–40 cm in length, 4–6 mm in width. Inflorescence erect, paniculate, 25–50 cm in length. Locally common on dry exposed sites at elevations of 120–1,920 m but notably common on dry exposed sites above 1,220 m on all of the main islands except Ni‘ihau, Kaua‘i, and Kaho‘olawe.

4.628 

Eragrostis variabilis

Erect perennial grasses with tufted culms, erect, 40–80 cm tall. Blades flat below, involute in their upper part, 50 cm in length, 10 mm in width. Inflorescence paniculate, up to 40 cm in length, reddish brown at maturity. Occurs on sand dunes, grasslands, open sites in dry

4.629 

Heteropogon contortus

FLOWERING PLANTS

Perennial grasses arising from tufted culms 40–100 cm tall, compressed. Blades bluish green, flat to slightly involute, 10–30 cm in length, 3–7 mm in width. Racemes solitary at the apex of flowering culms. Widely distributed throughout the tropics. Occurs on rocky cliffs, ledges, and on dry slopes near the coast to 700 m elevation on all of the main islands. Used by the Hawaiians as a source of thatching material because of its smell, color, and appearance. Thought to be indigenous but possibly a Polynesian introduction.

in width, margins distinctly thickened. Inflorescence an ovoid panicle, often purple, 10–20 cm in length, lower half or more of primary branches lacking spikelets. Occurs on wet ridgetops and in open places in wet forest at elevations of 215– 1,480 m on all of the main islands except Ni‘ihau and Kaho‘olawe. ENDEMIC | SECURE

4.632 

Isachne distichophylla Munro ex Hillebr. ‘ohe

ENDEMIC | SECURE

4.631 

Isachne pallens

Perennial grasses with decumbent culms, 30–60 cm tall, rooting at the nodes. Blades pale green, up to 6 cm in length, 2–3 mm in width, not marginally thickened. Inflorescence an ovoid panicle 3–8 cm in length. Occurs in mesic to wet forest, generally on rocks exposed to spray from waterfalls and along streams at elevations of 240–760 m on Kaua‘i and O‘ahu.

Isachne distichophylla

Erect perennial grasses with rather stout, stiff culms, decumbent at the base and rooting at lower nodes. Leaves distichous, green or often purple. Blades stiff and spreading, 5–17 cm in length, 4–10 mm

ENDEMIC | ENDANGERED

Isachne pallens Hillebr.

Isachne—A genus of annuals or herbaceous perennial grasses, often aquatic, with the culms ascending from a prostrate base and an open or contracted inflorescence. There are about 90 species found in the tropics and subtropics. The name is from the Greek iso, equal, and achne, scale, referring to the 2 similarly developed florets. In the Hawaiian Islands, there are 2 endemic species representing independent colonizing lineages.

4.630 

Ischaemum byrone (Trin.) Hitchc. Hilo ischaemum

Ischaemum—A genus of stoloniferous herbaceous perennial grasses with a terminal or axillary inflorescence often composed of paired racemes that are interlocked back to back. There are about 70 species found primarily in the paleotropics, particularly Asia. The name is from the Greek ischiamos, to quench blood flow, based on the use of the grass as a styptic for small cuts. Species exhibit C4 metabolism. In the Hawaiian Islands represented by 1 endemic and 1 naturalized species.

Ischaemum byrone

Perennial grasses with stolons and erect culms from a decumbent base, 40–80 cm tall. Blades flat, 7–20 cm in length, 3–5 mm in width. Inflorescences composed 2 or rarely 3 racemes, yellow to yellowish brown. Occurs among rocks near the ocean and sometimes farther inland at elevations up to 75 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i.

Lachnagrostis—A genus of about 20 species of largely perennial grasses with a center of distribution in the temperate Southern Hemisphere, especially Australasia. Closely related to Agrostis but separated on the basis of a hairy lemna and an elongated axis of the spikelet. Name comes from the Greek lachne, wooly, and agrostis, grass. In the Hawaiian Islands represented by the indigenous Lachnagrostis filiformis, a clump-forming perennial grass up to 60 cm tall. It occurs in a variety of habitats from dry ridges to the margins of bogs at elevations of 260–2,520 m on all of the main islands except Ni‘ihau. Listed in the Manual as Agrostis avenacea. Not pictured. Lepturus—A genus of herbaceous peren-

nial grasses with branched stolons, tufted culms, and a spicate, disarticulating inflorescence. About 10 species occur in coastal sites in the paleotropics from the east coasts of Africa to the Pacific Basin.

187

188

FLOWERING PLANTS

The name is from the Greek, leptos, slender, and oura, tail, referring to the slender inflorescences. Species exhibit C 4 metabolism. In the Hawaiian Islands represented by 1 indigenous species in the Northwestern Hawaiian Islands and Lehua.

Panicum beecheyi Hook. & Arn. panic grass

ENDEMIC | SECURE

Lepturus repens (G. Forst.) R. Br. INDIGENOUS | SECURE

4.634 

Panicum beecheyi

Tufted perennial grasses, culms unbranched, 30–60 cm in height. Blades 10–18 cm min length, 2–5 mm in width, flat or loosely inrolled. Inflorescence a panicle with spreading branches, 10–18 cm in length. Uncommon in dry, open, rocky habitats at elevations of about 900 m on Kaua‘i, O‘ahu, and Maui.

Panicum torridum Gaudich. kākonakona, panic grass ENDEMIC | SECURE

4.633 

Lepturus repens

Perennial grasses with branched stolons, culms erect or spreading, up to 60 cm tall, branching. Blades flat, stiff, up to 15 cm in length, 2–10 mm in width. Inflorescence a spike, 3–20 cm in length. Native from the Mascarene Islands across the Indian Ocean to Malesia, northern Australia and the Pacific Basin. Occurs on coastal sands just above the high tide mark on Kure, Midway, and Pearl and Hermes atolls, Lisianski, Laysan, French Frigate Shoals, and Lehua islet off Ni‘ihau. Often serves as an important stabilizing cover on sands.

Paspalum—A genus of annuals and rhizomatous or stoloniferous grasses with about 330 species. The group is pantropical but with a center of diversity in the neotropics. The name comes from the Greek paspalos, the name of a small millet mentioned in Greek literature. In the Hawaiian Islands represented by the indigenous Paspalum scrobiculatum, which occurs as a common colonist on slopes with thin soils and also in wet swampy ground at elevations from sea level to 1,070 m on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i. This species is possibly an inadvertent Polynesian introduction. Poa—A genus of annuals or sometimes stoloniferous or rhizomatous perennial grasses with an open to contracted inflorescence. There are at least 500 species that are found primarily in temperate and arctic regions worldwide. The name is from the Greek poa, a grass used as fodder for cattle. In the Hawaiian Islands represented by 3 endemic species from a single colonizing lineage and 2 naturalized species.

Poa sandvicensis (Reichardt) Hitchc. ENDEMIC | ENDANGERED

Panicum—A large genus of caespitose

annuals or herbaceous perennial grasses that are either rhizomatous or stoloniferous; the inflorescence always a panicle. About 300 species are known from warm and temperate areas worldwide. The name is from the old Latin word for millet. The genus includes both species with C3 and with C 4 metabolism. In the Hawaiian Islands, there are 14 endemic species representing 2 independent colonizing lineages and 6 naturalized species.

the main islands. The most common and widely distributed native Panicum in the Hawaiian Islands.

4.636  4.635 

Panicum torridum

Tufted annual grasses, culms branched or unbranched, 8–90 cm tall. Blades 4–29 cm in length, 2–15 mm in width, velvety hairy. Inflorescence a panicle with ascending branches, pubescent, 3–23 cm in length. Common in arid, weedy coastal flats at elevations from sea level to 500 m on Necker, Nīhoa, and all of

Poa sandvicensis

Tufted perennial grasses with short rhizomes and erect or slightly decumbent culms, 30–100 cm tall, slightly compressed laterally. Blades 10–20 cm in length, up to 6 mm in width. Inflorescence an ovoid panicle, 3–5 cm or more in length, branches whorled. Rare in wet and diverse mesic forest, especially shaded slopes, wet ridges, and ledges at elevations of 1,060–1,220 m on Kaua‘i.

FLOWERING PLANTS

Only 4 extant populations are known, with a total of about 6,000 individuals, with most of these in just 1 population. Primary threat comes from competition with alien plant species.

Poa siphonoglossa Hack. ENDEMIC | ENDANGERED

soaked, forcing out the seed. Species exhibit C 4 metabolism. In the Hawaiian Islands represented by 1 indigenous and 3 naturalized species.

Trisetum glomeratum (Kunth.) Trin. pili uka, mountain pili ENDEMIC | SECURE

Sporobolus virginicus (L.) Kunth

‘aki, ‘aki‘aki, mahiki, beach dropseed INDIGENOUS | SECURE

4.639 

4.637 

Poa siphonoglossa

Perennial grasses with short rhizomes and tufted culms, often extensive and cascading from banks in masses to 4 m in length, old culms naked and rushlike. Blades flat, typically less than 10 cm in length, 2–3 mm in width. Inflorescence an ovoid panicle, primary branches 2–3 cm in length, spikelets relatively few per panicle branch. Rare on wet, shady banks and along ridges at elevations of 480– 1,300 m in the Kōke‘e area on Kaua‘i. Recent surveys found 6 populations with about 50–70 clumps. Threats come from feral pigs, deer, and alien plant species.

Sporobolus—A genus of annuals or rhi-

zomatous perennial grasses with often tufted culms and an open, diffuse, or spikelike inflorescence. There are about 160 species that are cosmopolitan in temperate and tropical regions. The name is from the Greek sporos, seed, and ballein, to throw, referring to the structure of the fruit, which swells and bursts when

4.638 

Sporobolus virginicus

Perennial grasses spreading by extensive stiff rhizomes, culms erect, 5–50 cm or more in length. Blades involute, 3–14 cm in length, 2–5 mm in width. Inflorescence paniculate, dense, cylindrical, 2–9 cm in length, spikelets grayish or yellowish brown. Native to sandy sites usually near the coast in tropical and subtropical areas worldwide. Common on dunes and other coastal habitats from just above the high tide line to 15 m elevation on Midway Atoll, Laysan, and all of the main islands.

Trisetum—A genus of herbaceous perennial grasses with densely tufted culms and a narrow to open inflorescence. About 75 species are found in arctic and temperate regions worldwide and a few in montane tropical regions. The name is from the Latin tri, three, and seta, bristle, referring to the 3 bristles in each floret. In the Hawaiian Islands represented by 2 endemic and 1 naturalized species.

Trisetum glomeratum

Perennial grasses with densely tufted erect culms, 40–90 cm in length. Blades flat, coriaceous, 10–25 cm in length. Panicles erect, spikelike, sometimes loose or interrupted, 12–20 cm in length. Common on arid cinder fields, subalpine shrubland, and grassland and also in open bogs at elevations of 750–4,090 m on Maui and Hawai‘i.

Potamogetonaceae (PONDWEED FAMILY)

Freshwater aquatic herbs. Leaves floating or submerged. There are 7 genera and about 115 species with a nearly cosmopolitan distribution, with pondweed (Potamogeton) as the primary genus. In the Hawaiian Islands represented by the indigenous genus Potamogeton.

Potamogeton—A genus of perennial freshwater aquatic herbs with about 100 species and an almost cosmopolitan distribution centered in North America. The name comes from the Greek potamos, river, and geiton, neighbor, alluding to its aquatic habitat. In the Hawaiian Islands, there are 2 indigenous species representing independent colonizing lineages.

189

190

FLOWERING PLANTS

Potamogeton nodosus Poir. long-leaved pondweed

INDIGENOUS? | SECURE

worldwide with 1–7 species. The name honors Heinrich Ruppius, a seventeenth-century German botanist. Represented in the Hawaiian Islands by 1 indigenous species.

Ruppia maritima L.

ditchgrass, widgeon grass INDIGENOUS | SECURE

about 360 species widely distributed in tropical and subtropical regions, with a few species reaching into temperate regions. Economically the family is of no importance. A few species of Smilax are cultivated. Represented in the Hawaiian Islands by the indigenous genus Smilax.

Smilax—A genus of climbing shrubs and lianas. Plants are dioecious. There are about 350 species worldwide in tropical and temperate regions. The name is from the Greek smilax, the name for members of this genus. Represented in the Hawaiian Islands by 1 endemic species.

Smilax melastomifolia Sm.

pi‘oi, aka‘awa, hoi kuahiwi, catbrier ENDEMIC | SECURE

4.640 

Potamogeton nodosus

Plants only partially submerged, stems stout 40–150 cm in length, almost terete, arising from a dense mat of thick rhizomes. Leaves dimorphic, submerged ones thin and flaccid, linear to elliptic-lanceolate; floating leaves coriaceous, elliptic-linear, 1–10 cm in length, 10–20 nerved. Flowers numerous in congested spikes. Occurs infrequently at low elevations from sea level to 190 m in taro paddies, ditches, and ponds on O‘ahu and Maui. Questionably indigenous.

Ruppiaceae

(DITCHGRASS FAMILY)

Submerged aquatic perennial herbs. A single genus with about 7 species, although some systematists would lump all into a single variable species. Key features are the linear leaves with an expanded base consisting of the adherent stipules, inconspicuous flowers borne on short terminal spikes, and the small ovoid fruit borne on elongated slender stipes in an umbel-like cluster. Found along the coast but occasionally inland, although primarily plants of marine environments. Sometimes placed in the Potamogetonaceae.

Ruppia—A variable aquatic plant found in brackish, alkaline, or subsaline water

4.642  4.641 

Ruppia maritima

Submerged aquatic plants with very fine or slender, many-branched stems that are greenish to brownish. Leaves all similar, well spaced to somewhat clustered, 1–10 cm in length. Flowers axillary, 2 per peduncle, fruit ovoid, 2–3 mm in length. Nearly cosmopolitan plant of alkaline, subsaline, or brackish water. Occurs in brackish ponds, estuaries, and manmade channels near sea level on all of the main islands except Kaho‘olawe.

Smilacaceae

(CATBRIER FAMILY)

Climbing vines or lianas and occasionally perennial herbs or shrubs. Key features are the presence of creeping, starchy rhizomes, stems that are occasionally twining and often with recurved prickles, usually a pair of tendrils near the base of the leaf petioles, and often an umbellate inflorescence. There are 2 genera and

Smilax melastomifolia

Robust lianas arising from tuberous rhizomes, stems smooth or prickly, leaves coriaceous, heart shaped, 7–18 cm in length, with many conspicuous nerves. Plants dioecious. Flowers axillary in the upper leaves, 10–20 per umbel. The round berries are white at maturity. Occurs in mesic to wet forest at 180–2,080 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Xanthorrhoeaceae (GRASSTREE FAMILY)

Herbaceous perennials, leaf succulents, rosette succulents, and shrubby to tall semiwoody succulents and grasstrees. Widely distributed in the Southern Hemisphere and Eurasia, with 35 genera and about 900 species. Once considered an endemic Australian family with the single genus Xanthorrhoea, but recent molecular studies have now placed the Asphodelaceae and Hemerocallidaceae as subfamilies within a broader concept

FLOWERING PLANTS

of the Xanthorrhoeaceae. Species of Aloe are widely planted and naturalized easily. Represented in the Hawaiian Islands by 1 indigenous genus.

Dianella—A genus of rhizomatous perennial herbs with about 25 species found in the Mascarene Islands, tropical Asia, Australia, New Zealand, and on some Pacific islands. Formerly placed in a broadly defined Liliaceae and later in the Hemerocallidaceae. The name is from Diana, the Greek goddess of the hunt and the moon, and the Latin ella, the diminutive. The bright blue fruit characterizes this genus. In the Hawaiian Islands represented by 1 variable indigenous species. This family was included in the Liliaceae in the Manual.

Dianella sandwicensis Hook. & Arn. ‘uki, ‘uki‘uki

INDIGENOUS | SECURE

4.643 

Dianella sandwicensis

Coarse perennial herbs up to 2 m in height arising from rhizomes. Flower color varies from white to pale blue. Fruit color varies from dark translucent blue to sky blue. Occurs in the Marquesas and the Hawaiian Islands, where it is found in shaded sites in mesic to wet forest, dry shrubland, grassland, and lava fields at elevations of 120–2,040 m on all of the main islands except Ni‘ihau and Kaho‘olawe. A variable plant with respect to the inflorescence, flower and fruit color, and leaf width. The juice from the berries was used by the Hawaiians to dye kapa.

191

Cibotium chamissoi

CHAPTER 5

Ferns and Fern Allies

A

In this chapter, families of ferns and fern allies are presented alphabetically and genera and species alphabetically within their group. The scientific names presented here at the family, genus, and species level follow those used in the Smithsonian’s online Flora of the Hawaiian Islands, available at http://botany .si.edu/pacificislandbiodiversity/hawaiianf lora/index.htm. Refer­ence is made to names that have changed and family positions that have been rearranged compared to usage in Palmer (2003). The scientific name is followed by the Hawaiian name and an indication of the species as indigenous (i.e., occurs naturally in the Hawaiian Islands as well as elsewhere) or endemic (i.e., occurs only in the Hawaiian Islands). Endemism is indicated for the species level, and thus endemic varieties or subspecies are still noted as indigenous. Elevations given with species distributions follow those in Palmer. The conservation status of species as secure, rare, vulnerable, endangered, or extinct is also shown (see chapter 8 for definitions). Secure is used in the sense of not falling into one of the other categories and does not necessarily indicate that the species has large and/or numerous populations. Figures on the global number of species for individual families and genera generally follow those in Smith et al. (2006).

s described in chapter 2, the flora of ferns and fern allies in the Hawaiian Islands includes 163 native species, 2 more than listed in Palmer (2003), although with some rearrangements in systematic ranks. This species total represents 13% of the vascular plant flora. This proportion is high compared to the global proportion of ferns and fern allies, which represents about 4% of all vascular plants. However, this is not an unusually high proportion for tropical floras, which commonly have a rich diversity of ferns and fern relatives. Endemism is relatively high in Hawaiian ferns and fern allies by global standards, but it is lower than that of Hawaiian flowering plants. This is not surprising because of the ease of long-distance dispersal of tiny fern spores. Nevertheless, 71% of Hawaiian fern and fern allies are endemic, along with 2 endemic genera. These are Adenophorus (Polypodiaceae, formerly separated within the Grammitidaceae) and Sadleria (Blechnaceae). Despite this level of endemism at the species level, many of these lineages have not undergone extensive speciation after arrival in the Islands. Current estimates suggest that 104 founding lineages have produced the present flora of 164 species.

Aspleniaceae Terrestrial to epiphytic ferns. Key features are rhizomes creeping, ascending, or suberect, bearing net-form scales at apices and petiole bases. Sori linear and positioned along the veins and covered by a flap-shaped indusium arising along one edge. The family is widespread around the world but most diverse and common in tropical regions. Depending on individual views of systematists, there are anywhere from 1 to 7 genera and about 800 species, but 2 genera are recognized here. Several species of Asplenium, Phyllitis, and Camptosorus are grown as ornamentals. In the Hawaiian Islands represented by the indigenous genera Asplenium and Hymenasplenium. Diellia, formerly considered to be an endemic

genus of 6 species, has now been merged into Asplenium.

Asplenium acuminatum Hook. & Arn. lola ENDEMIC | SECURE

Asplenium—A large genus of about 650

species that are worldwide in distribution. They are terrestrial, epiphytic, or growing on rocks (rupestral or epilithic), with simple to compound fronds and short to elongate sori covered by an elongate indusium. The name is from the Greek a, without, and splen, spleen, referring to an old medicinal use. Several species are cultivated, such as Asplenium nidus, the bird’s-nest fern, which also is indigenous. In the Hawaiian Islands represented by 18 endemic and 9 indigenous species (2 of these represented by endemic subspecies), which are thought to represent 17 different colonizing lineages. Species formerly placed in the endemic genus Diellia are now placed here.

193

5.1 

Asplenium acuminatum

194

FERNS AND FERN ALLIES

Medium-sized to large ferns with large, dark green, and bipinnate fronds up to 200 cm in length; pinnules slightly lobed and marginally toothed. Locally common in mesic to wet forests at 275–2,500 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Aspleniun adiantum-nigrum L. ‘iwa‘iwa INDIGENOUS | SECURE

Medium-sized terrestrial ferns arising from a short rhizome. Fronds 55–130 cm in length; stipes typically densely covered with dark brown scaly hairs, giving rise to the specific epithet. Blades dark green, once pinnate with 24–48 pairs of closely set linear-lanceolate pinnae; sori close and parallel to the costae. Common in dry to mesic or wet forest at 200–1,000 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. Also occurs in Tahiti and Malesia. Listed in Palmer (2003) as Asplenium horridum.

Medium-sized terrestrial fern, rhizome erect. Fronds 20–95 cm in length, stipes 1/8th to 1/6th of the frond length. Blades once pinnate, narrow lanceolate, chartaceous, tips resemble lateral pinnae. Stipes and rachises dull brown, with conspicuous scales at the base of the stipes. Once locally common in mesic forests at elevations of 400–1,000 m on O‘ahu but now restricted to the Wai‘anae Mountains. Currently known from 13 populations totaling 4,000–7,000 individuals. Listed as Diellia falcata in Palmer (2003).

Asplenium contiguum Kaulf.

Asplenium diellaciniatum Viane

ENDEMIC | SECURE

5.2 

ENDEMIC | ENDANGERED

Asplenium adiantum-nigrum

Small terrestrial ferns from a stout, short horizontal rhizome. Fronds 10–40 cm in length; blades 2 to 3 pinnate with glossy pinnae. Stipes dark brown, exhibiting a distinct pattern of sharp color change to the light green of the lower rachises. Sori abundant in 2 rows next to the pinna midrib, ultimately joining to cover almost the entire lower surface of the pinnae. Locally common in open shrublands to dry woodlands from 350–4,000 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. On Hawai‘i and Maui growing on bare a‘ā lava flows in subalpine and alpine forest. Also found in Europe, Asia, and Africa.

Asplenium caudatum G. Forst. ‘iwa INDIGENOUS | SECURE

5.6 

5.4 

Asplenium contiguum

Small to medium-sized epiphytic ferns. Blades 1 pinnate, with sori in 2 rows parallel with the major vein. Common in mesic to wet forests at 335–1,675 m on all of the main islands except Ni‘ihau and Kaho‘olawe. There are 2 varieties, with the var. contiguum shown.

Asplenium diellaciniatum

Medium-sized terrestrial fern, sometimes growing over rocks. Fronds 20–70 cm in length, blades typically once pinnate; pinnae variable in shape. Rare in dry to mesic forests at elevations of about 1,000 m on Kaua‘i (Halemanu). Listed in Palmer (2003) as a synonym of Diellia erecta.

Asplenium dielpallidum N. Snow ENDEMIC | ENDANGERED

Asplenium dielfalcatum Viane ENDEMIC | ENDANGERED

5.7 

5.3 

Asplenium caudatum

5.5 

Asplenium dielfalcatum

Asplenium dielpallidum

Medium-sized terrestrial ferns. Fronds 20–40 cm in length, with the blade comprising 65–80% of the length; blades once pinnate. Stipes and rachis light brown in color. The species name comes from

FERNS AND FERN ALLIES

the pale green color of the blades. An extremely rare species known only from a few locations (Koai‘e, Ku‘ia, Mahanaloa, Mākaha) in mesic forests at 600–800 m elevation on Kaua‘i. Listed as Diellia pallida in Palmer (2003).

Asplenium lobulatum Mett. pi‘ipi lau manamana INDIGENOUS | SECURE

Asplenium nidus L. ‘ēkaha, ‘ēkaha kuahiwi

Asplenium hobdyi W. H. Wagner ENDEMIC | SECURE

5.8 

Fronds clustered at apex, 15–40 cm in length. Blades 1–3 pinnate, linear to linear-​lanceolate. Common in mesic to wet forests on all of the main islands except Ni‘ihau and Kaho‘olawe.

INDIGENOUS | SECURE

Asplenium hobdyi

Small terrestrial ferns. Fronds 10–35 cm in length, with blades once pinnate with 2–6 proliferations on the lower half of the blade. Uncommon and local in dark, moist places in wet forests at 1,200–1,850 m elevation on Kaua‘i, Moloka‘i, Maui, and Hawai‘i.

Asplenium kaulfussii Schldt. kūau ENDEMIC | SECURE

5.10 

Asplenium lobulatum

Medium-sized terrestrial ferns. Fronds 30–65 cm in length, blades coriaceous, once pinnate, varying from lobed to deeply cleft to the costae. The apex of the fronds typically with proliferations. Widespread in mesic to wet forest from 240–1,525 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Asplenium macraei Hook. & Grev. ‘iwa‘iwa, lau li‘i ENDEMIC | SECURE

5.12 

Asplenium nidus

Large epiphytic or terrestrial ferns with a bird-nest growth form. Rhizomes thick, short, and erect. Fronds simple, sessile, radiating from a center point, coriaceous, 50–100 cm or more in length. Common epiphyte in mesic to dry forest at elevations of 40–610 m all of the main islands except Ni‘ihau and Kaho‘olawe.

Asplenium normale D. Don INDIGENOUS | SECURE

5.9 

Asplenium kaulfussii

Medium-sized terrestrial ferns, occasionally epiphytic or growing over rocks. Fronds 25–85 cm in length, with blades typically once pinnate. Recognized by glossy, bright green, and succulent fronds that are prominently veined. Locally common in mesic to wet forests at 150– 1,200 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

5.11 

Asplenium macraei

Small to medium-sized terrestrial ferns. Extremely variable in morphology. Rhizomes trailing. Stipes dark brown with thin greenish ridges on the upper surface.

5.13 

Asplenium normale

Medium-sized terrestrial fern. Fronds 15–45 cm in length, blades once pinnate, narrow linear-lanceolate, dark green, coriaceous. With a single proliferation at

195

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FERNS AND FERN ALLIES

the tip of the rachis. Locally common in mesic to dry forest, rarely wet forest, from 375–1,680 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Asplenium peruvianum Desv.

INDIGNEOUS  |  ENDANGERED VARIETY

Medium-sized terrestrial ferns arising from a long, creeping rhizome. Fronds 22–130 cm in length, never proliferous; blades varying from once pinnate to fully 2 pinnate; 15–20 pairs of pinnae are sharply angled on the upper base. Stipes dark brown in color and covered with long, scaly hairs. Common in mesic forest on all of the main islands except Ni‘ihau and Kaho‘olawe.

Asplenium sphenotomum Hillebr. ENDEMIC | SECURE

5.14 

Hymenasplenium—A genus of about 30 species of epiphytic or terrestrial ferns, pantropical in distribution with a center of diversity in southwestern China. These species are separated from Asplenium on the basis of the peculiar vascular system in the dorsiventral rhizome. Long placed within Asplenium molecular studies show that Hymenasplenium represents a basally diverged lineage within the Aspleniaceae. In the Hawaiian Islands there is one indigenous species.

Asplenium peruvianum var. insulare

Delicate, small to medium-sized terrestrial fern. Fronds up to 50 cm in length. Often having proliferations on the stipe and midrib of the lower blade. Blades are once pinnate and linear in shape. Rare in dark, moist crevices and mouths of lava tubes at 1,650–2,200 m elevation on Maui and Hawai‘i. The Hawaiian plants represent the endemic var. insulare, with the typical form occurring in the Andes of South America.

Asplenium polyodon G. Forst. pūnana manu ENDEMIC | ENDANGERED

Hymenasplenium unilaterale (Lam.) Hayata pāmoho INDIGENOUS | SECURE

5.16 

Asplenium sphenotomum

Medium-sized terrestrial ferns, sometimes growing on mossy logs. Fronds 30–100 cm in length; blades typically 3 pinnate; proliferations present near the tips of the rachises. Locally common in wet forests at 1,140–1,970 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Asplenium trichomanes L. ‘oāli‘i INDIGENOUS | SECURE

5.15 

Asplenium polyodon

Small terrestrial ferns arising from a short thick caudex. Fronds 7–30 cm in length; blades once pinnate with 15–45 round to ovate pinnae on each side; lowest pinnae small and distantly arranged. Stipes densely tufted, wiry and shiny dark brown, with old stipe bases retained on the rhizome. Locally abundant in subalpine and alpine areas at 1,200–2,700 m elevation on East Maui and Hawai‘i. Virtually cosmopolitan in temperate regions of the world and on high mountain regions of the tropics. Hawaiian plants are the subsp. densum, which is endemic.

5.17 

Asplenium trichomanes var. densum

5.18 

Hymenasplenium unilaterale

Small to medium-sized terrestrial ferns, occasionally growing over rocks. Rhizomes long and creeping. Fronds 15–35 cm in length; blades once pinnate, with their tips tapering gradually and evenly. Locally common in wet mossy forests, particularly near streams, at 200–1,830 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Athyriaceae—This

family name has now been changed to the Woodsiaceae.

FERNS AND FERN ALLIES

Blechnaceae

Small to large terrestrial, sometimes climbing ferns. Key features are the presence of anthocyanins in the unfolding leaves that color them reddish, sori in chains or linear, often parallel and adjacent to midribs, and linear indusia opening inwardly toward midribs. Petioles with numerous round vascular bundles arranged in a ring. There are 9 genera and about 200 species that are cosmopolitan in distribution. Several species of Blechnum are cultivated as ornamentals. Represented in the Hawaiian Islands by the endemic genus Sadleria and the indigenous genus Doodia.

Doodia—A genus of 17 species of small to medium-sized terrestrial ferns distributed across the Indian and Pacific oceans from Sri Lanka to the Hawaiian Islands and Rapa Nui. The tufted fronds are pinnate or pinnatifid, with numerous sori per segment. The genus honors Samuel Doody, a London apothecary of the nineteenth century. In the Hawaiian Islands, there are 2 endemic species. An interspecific hybrid has been collected.

green lanceolate pinnae, merging together above and separated below. Sori in parallel rows on each side of the costae. Common in mesic or sometimes dry forests from 90–1,220 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Sadleria—A genus of 6 species of mostly terrestrial ferns endemic to the Hawaiian Islands. The sori borne in 2 parallel lines on each side of the segment midrib and the presence of a mass of soft, brownish scales at the bases of the frond stems are characteristic. Species resemble a small tree fern, with heights up to 1.5 m and a trunk diameter of up to 10 cm. The genus honors Joseph Sadler, a nineteenth-century Hungarian physician who studied ferns. The Native Hawaiians used the hairs of this fern (pulu) for stuffing pillows and other objects. The stipes were used as sizing in making kapa, while the fronds were often used as a mulch. In hard times, the pith of the trunk was cooked and eaten.

Sadleria cyatheoides Kaulf. ‘ama‘u, ma‘u, ma‘uma‘u ENDEMIC | SECURE

Doodia kunthiana Gaudich. ōkupukupu, pāmoho, ōkupukupu lau‘i‘i ENDEMIC SECURE

pairs of segments on each pinnae; blades often reddish when young. The glabrous, straw-colored stipes have a mass of soft brown scales at their base. Common in mesic, wet, or even subalpine forests at 75–2,200 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Sadleria pallida Hook. & Arn. ‘ama‘u, ma‘u, ma‘uma‘u ENDEMIC | SECURE

5.22 

Sadleria pallida

Medium-sized terrestrial ferns with long, branching rhizomes. Fronds erect, up to 100 cm or more in length; blades once pinnate-pinnatifid to twice pinnate. A distinguishing feature from other species is that ultimate segments on each pinna are broad and usually less than 30 in number. Common in wet forests and on windswept rainy ridges at 25–2,150 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Sadleria souleyetiana (Gaudich.) T. Moore ENDEMIC | SECURE

5.19 

5.20 

Sadleria cyatheoides—habit

5.21 

Sadleria cyatheoides

Doodia kunthiana

Medium-sized terrestrial ferns arising from a short, creeping rhizome. Fronds 25–65 cm or more in length and arranged in rosettes; stipes about half of the frond length; blades once pinnate with dark

Terrestrial ferns with trunks from 1.0 to 1.5 m tall. The smooth, erect, and coriaceous fronds are up to 300 cm in length; blades bipinnate with 30–60 dark green

5.23 

Sadleria souleyetiana

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FERNS AND FERN ALLIES

Medium-sized to large terrestrial ferns. Fronds held erect and 50–350 cm in length; blades once pinnate to 2 pinnate near the base of large fronds. A distinctive feature is the presence of mats of scales resembling wet tissue paper at the base of the stipes. Occasional in mesic to wet forest at 400–1,550 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Sadleria squarrosa (Gaudich.) T. Moore ‘apu‘u ENDEMIC | SECURE

Cibotiaceae

Large terrestrial ferns, decumbent to erect and arborescent. Key features are long-matted trichomes at the base of the petioles and many fibrous roots that can occur from the base to the apex of the stem. The family includes a single genus distributed in tropical mountain thickets and forests of Southeast Asia (4 species), Mexico and Central America (2 species), and the Hawaiian Islands. Represented in the Hawaiian Islands by Cibotium.

Cibotium—A genus of 11 species of large terrestrial ferns as described above. The name is from the Greek kibotos, box, referring to the shape of the indusium. Some species are cultivated. In the Hawaiian Islands, there are 4 endemic species and 3 interspecific hybrids.

Tree ferns 1–2 m and occasionally up to 6 m in height. Fronds up to 5 m long. The undersides of the blades are light green in color and clothed with tan, cobwebby hairs. Distinctive features are stipes naked except for golden or mustard yellow hairs near their base and a skirt of dead fronds. In mesic to wet forests at 150–1,200 m elevation (less common above 800 m) on all of the main islands except Kaua‘i, Ni‘ihau, and Kaho‘olawe.

Cibotium glaucum (Sm.) Hook. & Arn. hāpu‘u, hāpu‘u pulu ENDEMIC | SECURE

Cibotium chamissoi Kaulf. ENDEMIC | SECURE

5.24 

Sadleria squarrosa

5.25 

Cibotium chamissoi—habit

Small to medium-sized terrestrial ferns with a creeping rhizome. Fronds drooping, 15–32 cm in length; blades 2 pinnate, coriaceous in texture; pinnae often lobed and rounded, giving a beaded appearance; stipes and rachises heavily clothed with scales. Infrequent in wet forest on wet or damp cliff faces or slopes on all of the main islands except Ni‘ihau and Kaho‘olawe.

5.27 

Sadleria unisora (Baker) W. J. Rob. ‘apu‘u ENDEMIC | SECURE

Small terrestrial ferns. Fronds drooping, up to about 40 cm in length. Similar to S. squarrosa but with narrower stipes and rachises and round-tipped pinnae. Uncommon along stream walls in wet forests of Kaua‘i at 400–1,520 m elevation (see figure 2.6).

5.26 

Cibotium chamissoi

Cibotium glaucum

Tree ferns 1–2 m and occasionally to 6 m tall. Fronds to 7 m in length. The characteristic glaucous blue gray undersurface with white cobwebby hairs separates this species from others. The stipes are clothed at the base with fawn-colored matted hairs. A common species of mesic and wet forest at 300–1,700 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. Occasionally hybridizing with C. menziesii.

FERNS AND FERN ALLIES

Cibotium menziesii Hook. hāpu‘u i‘i ENDEMIC | SECURE

lobes forming a scalelike false indusium. In the Hawaiian Islands represented by 1 endemic species.

Microlepia strigosa (Thunb.) Presl. palai, palapalai INDIGENOUS | SECURE/RARE

Hypolepis hawaiiensis Brownsey olua ENDEMIC | SECURE

5.28 

Cibotium menziesii—habit

Tree ferns usually 2–7 m tall. Fronds up to 5 m in length. Distinguished by stipes covered with stiff, dark brown hairs and a dark green upper surface and an absence of cobwebby hairs on the lower surface of the blades. Common in wet to mesic forests at elevations of 250–1,400 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Dennstaedtiaceae

Terrestrial ferns. Key features are the long, creeping stems that bear large decompound leaves, the presence of marginal or submarginal sori, and in some species buds on the petioles developing into branches. The indusia are linear or cup shaped and positioned at the blade margin or reflexed over the sori. There are 11 genera and about 170 species found mostly in the tropics but a few extending into temperate or boreal regions. Some genera, such as Pteridium, Microlepia, and Hypolepis, are cultivated. Young fiddleheads of Pteridium are gathered as a food source. In the Hawaiian Islands represented by the indigenous genera Hypolepis, Microlepia, and Pteridium.

Hypolepis—A genus of about 55 tropical

and subtropical species of medium to large terrestrial ferns. Sori lack indusia and are sometimes covered by the partially curled margins of the pinnae. The Hawaiian taxa are large terrestrial ferns superficially resembling Pteridium aquilinum, but they differ in having round, single sori placed marginally or submarginally compared to the linear and often continuous sori of Pteridium. The name comes from the Latin hypo, below, and lepis, scale, referring to the reflexed leaf

5.30  5.29 

Hypolepis hawaiiensis

Small to large-sized terrestrial ferns. Fronds 25–120 cm in length; blades 2 to 4 pinnate with an ovate to broadly ovate shape; sori submarginal and sometimes partially covered by the margin of the pinnae. Resembles Pteridium aquilinum but differs in having individual round sori, neither continuous nor linear. Locally common in wet forests at 600–1,980 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. There are 2 varieties, with the typical form pictured. The diminutive var. mauiensis is known only from Pu‘ukukui and ‘Eke on West Maui.

Microlepia—A genus of about 70 species of

mostly paleotropical terrestrial ferns but extending from Japan to New Zealand. Often occurring in moist to wet forests, sometimes dry forests, and along roadsides and open places. The genus is characterized by long, creeping rhizomes, blades 2–3 pinnate, and abaxial to marginal sori with a half cup-shaped indusium. The name is from the Greek mikros, little, and lepis, scale, in reference to its small cup-shaped indusia. Some species are cultivated as ornamentals and used in lei and hula altars. Represented in the Hawaiian Islands by 2 indigenous species and 1 natural hybrid.

Microlepia strigosa var. strigosa

Medium-sized terrestrial ferns. Fronds usually less than 100 cm in length but occasionally 300 cm long; blades bipinnate-pinnatifid to 3 pinnate-pinnatifid, with pale-colored and prominent raised veins on the lower surface; sori marginal to submarginal and covered by an indusium attached at both the base and sides. Occurs in dry to mesic forest from sea level to 1,770 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. There are 2 varieties present, with the typical form shown here. The rare var. mauiensis is endemic to Maui and Hawai‘i. This fern was gathered by the Hawaiians and used in the making of the hula altar honoring Laka, the goddess of the hula.

Pteridium—A genus of 5 species cosmopolitan in distribution, sometimes considered as a single polymorphic species. Known widely as bracken fern, it grows in a wide variety of habitats, often becoming weedy. The genus is characterized by wide creeping, multibranched stems. The plant sends up large, triangular fronds from a creeping underground rootstock and may form dense thickets. This rootstock often extends 1 m or more underground between fronds. The name is from the Greek diminutive of pteris, the ancient name for ferns. Although young fiddleheads of this fern are sometimes gathered

199

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FERNS AND FERN ALLIES

as a food source, they contain a nerve poison and are also toxic to livestock. The plant is used for thatch, bedding material for animals, and packing material for fruits.

Pteridium aquilinum (L.) Kuhn kīlau, bracken fern INDIGENOUS | SECURE

in form but are absent in some taxa. The family is cosmopolitan in distribution, but the number of genera differs according to individual authors, especially in eastern Asia where the family is most diverse. There are 40–45 genera and about 1,700 species, with the great majority of these species in just 4 genera (Ctenitis, Dryopteris, Elaphoglossum, and Polystichum). Many species of Cyrtomium, Dryopteris, and Polystichum are cultivated. In the Hawaiian Islands represented by 5 indigenous genera: Arachniodes, Ctenitis, Cyrtomium, Dryopteris, and Polystichum.

Arachniodes—A genus of about 140 tropical and subtropical species of moderate to large terrestrial ferns, with a range from India across Southeast Asia and Australia to the Pacific and neotropics. The name comes from the Greek aracnion, spider’s web, and odes, having the form of, although it is not clear to what morphology this name refers. In the Hawaiian Islands represented by 1 endemic species.

stipes form half of the frond length, with a base covered in reddish brown scales. Infrequent but common in local areas along streams in wet forests at 765–2,015 m elevation on all of the main islands except Ni‘ihau, Lāna‘i, and Kaho‘olawe. Rare on O‘ahu and known only from the Kohala Mountains on Hawai‘i.

Ctenitis—A genus of about 120 species of terrestrial or rock-dwelling ferns found throughout the tropical and warm temperate regions of the world. The genus is characterized by the raised midrib on the upper surface of the segments and the presence of short, multicellular trichomes. The name is from the Greek ktenos, comb, referring to comb-shaped projections of the spores. In the Hawaiian Islands represented by 2 endemic species.

Ctenitis latifrons (Brack.) Copel. ‘ākōlea ENDEMIC | SECURE

Arachniodes insularis W. H. Wagner ENDEMIC | SECURE

5.31 

Pteridium aquilinum subsp. decompositum

Moderate-sized to large terrestrial ferns arising from a subterraean branching rhizome. Fronds 50–200 cm in length; blades 3 pinnate below, simply pinnate above, terminating in a pinnatified apex. Stipes with horseshoe-shaped vascular bundles in cross section. Common in mesic to wet shrublands and forests, grasslands, and subalpine habitats at elevations of 300– 2,700 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Hawaiian plants represent the endemic subsp. decompositum. Listed in Palmer (2003) as Pteridium aquilinum var. decompositum.

5.33 

Medium-sized to large terrestrial ferns. Fronds up to 150 cm in length; dark green blades 2 to 3 pinnate and ovate to deltoid in shape. Stipes covered with short overlapping hairs and accounting for about half of the length; lower third of stipe length, densely covered with lanceolate brown scales. Sori medial to marginal and indusia variably present. Occurs in mesic to wet forest and in particular in the understory of ‘ōhi‘a forests at 400–1,500 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Dicksoniaceae—The genus Cibotium was formerly placed in this family but has now been separated as the Cibotiaceae. Dryopteridaceae

Terrestrial ferns. The sori are generally round in shape and borne on the underside of the blades. Indusia are often peltate

Ctenitis latifrons

5.32 

Arachnoides insularis

Medium-sized to large terrestrial ferns. Fronds up to 130 cm in length; blades 3 to 4 pinnate and triangular-ovate in shape;

Cyrtomium—A genus of terrestrial or rock-growing ferns found in temperate and tropical regions of the world. Most species have an odd-pinnate blade with the margins of the pinnae usually bearing

FERNS AND FERN ALLIES

prominent teeth. There are about 25 species, with 15 or more found in East Asia. This genus is sometimes lumped into Polystichum. The name is from the Greek kyrtos, bent or curved, referring to the arc-shaped union of the veins. The holly fern, C. falcatum, is commonly cultivated and also naturalized in the Islands. In the Hawaiian Islands represented by 1 indigenous species.

Dryopteris crinalis (Hook. & Arn.) C. Chr. palalalai ‘aumakua ENDEMIC | SECURE/RARE

the pinnate fronds. Occurs in dry to wet forest at elevations of 500–2,100 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Two varieties are present, with the typical form illustrated here. The var. lamoureuxii is rare on East Maui.

Dryopteris glabra (Brack.) Kuntze kīlau, hohiu ENDEMIC | SECURE/RARE

Cyrtomium caryotideum (Wall.) C. Presl kā ‘ape‘ape, ‘ahina kuahiwi INDIGENOUS | SECURE

5.37  5.35 

5.34 

Cyrtomium caryotideum

Medium-sized terrestrial ferns arising from a large prostrate rootstock covered with the remains of old stipe bases. Fronds erect to spreading, 25–50 cm in length; blades once pinnate with 3–6 pairs of pinnae that are toothed and sometimes lobed, pale green (not glossy), and somewhat coriaceous. The sori are scattered all over the undersurface of the blade. Found widely in summer-dry mesic and wet forests and mesic shrublands on all of the main islands except Ni‘ihau and Kaho‘olawe. Also occurs in Asia.

Dryopteris crinalis var. crinalis

Medium-sized terrestrial ferns. Fronds 20–85 cm in length; blades 2 to 3 pinnate, lacking indusia. Distinguished by narrow and twisted dark brown scales and hairs on the stipes, rachises, and underside of the pinnae. Locally common in wet forests at 300–1,900 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. There are 2 varieties, with the typical form pictured. The var. podosorus is rare on Kaua‘i (Kōke‘e, Kawaikōī).

Dryopteris glabra var. glabra

Medium-sized terrestrial ferns. Fronds erect to spreading up to 100 cm in length and 40 cm in width. Distinguished by 3 to 4 pinnate blades and marginal to submarginal sori with indusia. A variable species with 7 named varieties. The form shown here is var. glabra. Common in mesic to wet forests at 460–2,200 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Dryopteris hawaiiensis (Hillebr.) W. J. Rob. ENDEMIC | SECURE

Dryopteris fusco-atra (Hillebr.) W. J. Rob. Ī‘i

ENDEMIC | SECURE/RARE

Dryopteris—A genus of terrestrial or

rock-growing ferns that are found in the temperate and tropical regions of the world. There are about 150 species, with the greatest diversity in Eastern Asia. The fronds are usually borne in a crown or cluster, with the blades usually pinnate to bipinnate or more. Most species have a kidney-shaped indusium. The name is from the Greek drys, oak, and pteris, fern. Many species are in cultivation. In the Hawaiian Islands represented by 10 endemic and 1 indigenous species, representing 6 independent colonizing lineages.

5.38 

5.36 

Dryopteris fusco-atra var. fusco-atra

Terrestrial ferns arising from an erect caudex. Fronds 30–80 cm in length, blades once pinnate with sori present on all pinnae; stipes dark in color and covered with pale brown scales below, becoming blackish along the rachis of

Dryopteris hawaiiensis

Terrestrial ferns arising from an erect caudex. Fronds up to 60 cm in length; blades 2 to 3 pinnate; stipes slender and sparingly covered throughout with thin brown scales changing to dark fibrils on the rachis of the fronds. Infrequent but locally common in mesic to wet forest at 600–2,000 m elevation on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i.

201

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FERNS AND FERN ALLIES

Dryopteris rubiginosa (Brack.) Kuntze ENDEMIC | SECURE

Dryoptris unidentata (Hook. & Arn.) C. Chr. ‘akole ENDEMIC | SECURE

5.39 

Dryopteris rubiginosa

Terrestrial ferns arising from a thick rhizome. Fronds to 100 cm in length; dark green blades 3 to 4 pinnate and deltoid in shape; stipe widely curved at the base with dark brown, curly scales below. Locally common in wet forest at elevations of 885–2,180 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in Palmer (2003) as Nothoperanema rubiginosum.

Dryopteris tetrapinnata W. H. Wagner & Hobdy ENDEMIC | ENDANGERED

5.41 

Dryopteris unidentata var. palacea

Large terrestrial ferns. Fronds 100–200 cm in length; blades 3 to 4 pinnate. Sori marginal and lacking indusia. Locally common in wet forests at 460–1,750 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. There are 2 named varieties, with the var. paleacea shown.

from the rhizome at a 45-degree angle; blades mostly once pinnate. Stipes and rachises heavily covered with reddish brown to brown scales. Occurs in mesic forest and open areas at 850–2,470 m elevation on all of the main islands except Ni‘ihau, Lāna‘i, and Kaho‘olawe.

Elaphoglossum—A genus of terrestrial, rupestral, or epiphytic ferns that are pantropic and south-temperate in distribution. There are about 600 species, with the major portion of these in the neotropics. The fronds have entire margins and are usually dimorphic, with fertile blades differing in size and shape. Lower surface of fertile blades covered with sporangia. The name comes from the Greek elaphos, deer, and glossa, tongue, referring to the shape of the fronds. In Hawai‘i, there are 8 endemic species, 2 endemic hybrids, and 1 indigenous species. This genus was formerly placed in the Lomariopsidaceae.

Elaphoglossum aemulum (Kaulf.) Brack. ‘ēkaha, ‘opeha, laukahi nunui ENDEMIC | SECURE

Dryopteris wallichiana (Spreng.) Hyl. laukahi INDIGENOUS | SECURE

5.43 

5.40 

Dryopteris tetrapinnata

Very large and showy terrestrial ferns. Fronds up to 300 cm in length; blades 4 pinnate and highly dissected. Sori lack indusia. Scattered in occurrence in the understory of wet forest at 1,400–1,830 m elevation on East Maui.

5.42 

Dryopteris wallichiana

Medium-sized to large terrestrial ferns. Fronds up to 150 cm in length with a shuttlecock arrangement that radiates

Elaphoglossum aemulum

Medium-sized epiphytic ferns, occasionally terrestrial. Plants growing on tree trunks often produce a broad mat of rhizomes, rhizome scales, and organic matter. Sterile blades 20–45 cm in length, coriaceous; veins parallel and free. Common in wet forests at 300–1,400 m elevation on all of the main islands except Ni‘ihau, Kaho‘olawe, and Hawai‘i (extinct).

FERNS AND FERN ALLIES

Elaphoglossum alatum Gaudich. hoe a Māui ENDEMIC | SECURE

Medium-sized epiphytic ferns arising from a prostrate, creeping rootstock covered with dense brownish scales. Sterile fronds 17–45 cm in length, strongly coriaceous in texture. It is the most common Hawaiian Elaphoglossum and the only species with a netlike vein pattern. Forms large populations in open mesic and wet forest at 300–1,300 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Elaphoglossum paleaceum (Hook. & Grev.) Sledge māku‘e, ‘ēkaha INDIGENOUS | SECURE

Elaphoglossum fauriei Copel. hoe a Māui, ‘ēkaha ENDEMIC | SECURE

5.44 

Elaphoglossum alatum

Medium-sized terrestrial ferns, occasionally epiphytic. Rhizomes short creeping with tangled masses of long brown scales; veins parallel, uniting near the margin to form a continuous marginal vein. Sterile blades 35–60 cm in length; stipes winged almost to their base. Common and sometimes found in large patches in shaded mesic to wet forests at 450–1,100 m elevation on O‘ahu, particularly in the Wai‘anae Mountains.

5.47 

Medium-sized epiphytic ferns, or occasionally terrestrial, arising from a short, thick, woody, creeping rootstock. Stipes winged, thickly clothed with dark scales. Sterile fronds 17–45 cm in length, covered on both sides with brownish scales, and often nodding in position rather than held upright. Common in mesic to wet forest at elevations of 875–2,075 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Elaphoglossum crassifolium (Gaudich.) W. R. Anderson & Crosby hoe a Māui, ‘ēkaha

Elaphoglossum pellucidum Gaudich.

ENDEMIC | SECURE

ENDEMIC | SECURE

5.46 

5.45 

Elaphoglossum crassifolium

Elaphoglossum paleaceum

Elaphoglossum fauriei

Medium-sized terrestrial or epiphytic fern with short, creeping rhizome. It is one of 4 Hawaiian species forming a group distinct from those others illustrated here, characterized by winged stipes and parallel and forking veins that combine to form a continuous marginal vein on the blade. Occurs in mesic to wet forests at elevations from 600–1,350 m in the Ko‘olau Mountains on O‘ahu and Moloka‘i.

5.48 

Elaphoglossum pellucidum

203

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FERNS AND FERN ALLIES

Small to medium-sized terrestrial or epiphytic ferns with long creeping rhizomes that often are vinelike and wind around tree trunks or rocks. Sterile blades 6–25 cm in length; thin and translucent, with both sides covered with shining glands, giving living plants a sticky surface. Uncommon in mesic forest at 450–1,270 m on all of the main islands except Ni‘ihau, Lāna‘i, and Kaho‘olawe.

indusium is usually peltate. The name is from the Greek polys, many, and stichos, rows, referring to the numerous regular rows of sori. Many species are cultivated for their attractive fronds. In the Hawaiian Islands, there are 3 endemic species.

Polystichum bonseyi W. H. Wagner & Hobdy ENDEMIC | SECURE

Elaphoglossum wawrae (Luerss.) C. Chr. ‘ēkaha

Gleicheniaceae

ENDEMIC | SECURE

5.50 

5.49 

Elaphoglossum wawrae

Small to medium-sized epiphytic ferns, occasionally terrestrial. Rhizomes short creeping, arising from a thick, woody, creeping rhizome. Sterile and fertile fronds almost equal in size and shape, with the former 7–25 cm in length, linear-lanceolate in shape, pale green, and leathery. While resembling E. crassifolium, it can be distinguished by its smaller size, narrower blades, and free and forked veins. Common in mesic to wet forests at elevations of 800–2,100 m on all of the main islands except Ni‘ihau, Lāna‘i, and Kaho‘olawe.

Medium-sized terrestrial ferns arising from a thick, erect rhizome with numerous fibrous roots. Fronds 45–95 cm in length; blades coriaceous, dark green and glossy, 45–95 cm in length; rachises densely covered with linear hairlike scales; stipes densely covered with large overlapping scales. Locally common in open areas and high-elevation wet forests at 1,400– 2,060 m elevation on Maui and Hawai‘i.

Polystichum bonseyi

Medium-sized terrestrial ferns. Rhizomes short creeping. Fronds up to 100 cm in length; blades bipinnate, lanceolate in shape. The distinguishing features are chartaceous blades that are only minimally or not at all narrowed at their base and stipes covered with brownish, hairshaped scales. Occurs in mesic and wet forests and subalpine wet shrublands at 1,400–2,000 m elevation on East Maui and known from a single collection from Hualālai on Hawai‘i.

Polystichum hillebrandii Carruth. ka‘upu, papa‘oi ENDEMIC | SECURE

Terrestrial ferns often sprawling over the ground and other vegetation. Key features are long creeping rhizomes and the pseudodichotomous fronds usually branching several times due to the arrested growth of the main divisions that develop in succeeding seasons. The sori are not marginal and lack an indusium. The family is comprised of 6 genera and about 125 species that are found primarily in the tropics or the subtropics of the Southern Hemisphere. Many species are colonizing invasives on disturbed sites and road cuts and are often weedy by nature. None are widely grown as ornaments. Represented in the Hawaiian Islands by 3 indigenous genera: Dicranopteris, Diploptergium, and Sticherus.

Dicranopteris—A pantropical genus of 20

species of terrestrial ferns. The fronds are large and pseudodichotomously branched. The stems and blades are covered with trichomes. The name is from the Greek dikros, forked, and pteris, fern, referring to the shape of the fronds. In the Hawaiian Islands, there is 1 indigenous species.

Dicranopteris linearis (N. L. Burm.) Underw. uluhe, unuhe INDIGENOUS | SECURE

Nothoperanema—This genus is now placed within Dryopteris.

Polystichum—A genus of terrestrial or rock-growing ferns that are cosmopolitan in distribution. There are close to 160 species. The simple to 3-pinnate-­ pinnatifid fronds are usually borne in a crown, with the petioles usually persistently scaly, especially at the base. The

5.51 

Polystichum hillebrandii

5.52 

Dicranopteris linearis

FERNS AND FERN ALLIES

Large terrestrial ferns with dichotomously branching fronds up to 1 m or more in height when supported by other vegetation. Often colonizing landslides and disturbed areas, forming extensive impenetrable thickets over bare ground or on top of other plants. The sori are borne in a row on each side of the costae. Rhizomes and stipes with a dominance of hairs rather than scales. Very common in open areas of mesic to wet forest from sea level to above 2,000 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. Native Hawaiians brewed an infusion from the fronds to serve as a laxative.

to bipinnate, with the rachis often pinnatifid in part. The name is from the Greek stichos, in a row, referring to the 2 rows of sori per pinna. In the Hawaiian Islands, there is 1 endemic species.

Sticherus owhyhensis (Hook.) Ching uluhe, unuhe ENDEMIC | SECURE

Callistopteris—A small genus of 5 epiphytic species ranging from Southeast Asia to the Pacific Islands. These species would fit within a broad concept of Trichomanes. The name comes from the Greek kallistos, most beautiful, and pteris, fern. In the Hawaiian Islands represented by 1 endemic species.

Diplopterygium—A predominantly Male-

sian genus of about 25 species of terrestrial ferns. The fronds are large, pinnate with bipinnate pinnae and an unbranched rachis, and exhibit indeterminate growth from dormant buds between the terminal pinnae of each blade. The name is from the Greek diploos, twofold, and pteris, fern, referring to the shape of the fronds. In the Hawaiian Islands, there is 1 indigenous species.

Diplopterygium pinnatum (Kunze) Nakai uluhe lau nui INDIGENOUS | SECURE

5.53 

Diplopterygium pinnatum

Medium-sized to large terrestrial ferns with creeping rhizomes. Fronds large, opposite; blades bipinnate and covered with brittle, often recurved, shiny dark scales, rachises unforked; stipes up to 40 cm in length. Occurs in wet forest, often on steep banks or streamsides at 350–1,500 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. Also in Japan, China, Malesia, and on other Pacific islands.

Sticherus—A pantropical and Southern Hemisphere genus of about 80 species of terrestrial ferns. The fronds are pseudodi­ chotomously branched, usually pinnate

shape. Recent studies have confirmed the monophylly of these 2 groups, but there are major disagreements among authors about natural generic groups within these 2 lineages. Up to 30 genera have been described, but an intermediate view recognizes 9 genera and about 600 species. A few species are cultivated as ornamentals. In the Hawaiian Islands, there are 4 indigenous genera: Callistopteris, Credidomanes, Hymenophyllum, and Vandenboschia. Palmer (2003) used a different grouping of genera for these species.

Callistopteris baldwinii (D. C. Eaton) Copel. ENDEMIC | SECURE

5.54 

Sticherus owhyhensis

Large terrestrial fern with creeping rhizomes resembling Dicranopteris in growth form but differing in that all or most of the branches bear pinnae and veins forked just once. Rhizomes and stipes with a dominance of scales rather than hairs. The sori are in one row on each side of the pinna midrib near the margin. Occurs sporadically in wet forest forming tangled masses at elevations of 900–1,740 m on all of the main islands except Ni‘ihau and Kaho‘olawe. 5.55 

Grammitidaceae—This family is now placed within the Polypodiaceae. Hymenophyllaceae

Primarily epiphytic ferns of tropical and wet temperate regions. Commonly known as filmy ferns. Key features are the thin blades 1 to a few cells thick (sometimes thalloid) that lack stomata. The veins are dichotomously branched and the sporangia enveloped in a cup-shaped indusium. The family formerly comprised 2 genera, Hymenophyllum and Trichomanes, separated on the basis of characters of sorus

Callistopteris baldwinii

Small to medium-sized terrestrial filmy ferns. Fronds clustered at the tip of erect rhizomes; blades up to 50 cm in length, deeply cut 3 to 4 pinnate; stipes conspicuously hairy. Highly variable in size depending on substrate. The small ribbon-shaped gametophytes of this species are reported to be more common than the sporophytes and often occur on wet banks in the absence of sporophytes. Locally common on soil banks in wet mossy forests at elevations of 300– 1,600 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

205

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FERNS AND FERN ALLIES

Crepidomanes—A genus of 35 epiphytic species, largely distributed in the paleotropics but with a few species in the neotropics. These species would fit within a broad concept of Trichomanes. In the Hawaiian Islands represented by 1 endemic and 1 indigenous species.

Crepidomanes draytonianum (Brack.) Copel. ENDEMIC | SECURE

Small filmy fern with creeping filamentous rhizomes. Fronds 2–6 cm in length, sometimes mat forming and appearing mosslike. This is the smallest Hawaiian fern and is easily overlooked when growing with mosses. Common on mossy rocks and trees in mesic to wet forests at elevations of 90–1,280 m on all of the main islands except Ni‘ihau and Kaho‘olawe. This species includes both Gonocormus minutus and G. proliferus, as described in Palmer (2003).

Hymenophyllum—A genus of about 275 species of epiphytic or epilithic ferns that are pantropical and south temperate in distribution, with a few species distributed in northern temperate areas. The fronds are thin, glabrous, and pinnately compound with a 2-valved indusium that has the receptacle included. The derivation of the name is from the Latin hymeno, membrane, and phyllum, leaf. In the Hawaiian Islands, there are 3 endemic species.

Hymenophyllum lanceolatum Hook. & Arn. palai hinahina ENDEMIC | SECURE

5.56 

Crepidomanes draytonianum

Small filmy fern with creeping filamentous rhizomes. Fronds spreading; blades 15–57 cm in length. Indusia cone shaped with a broad flaring tip. Uncommon but sometimes locally abundant in shady wet forests on moist rocks, stream banks, and mossy tree trunks. Broadly resembles a miniature Vandenboschia davalloides. Present in wet forests at elevations of 20–1,100 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in Palmer (2003) as Vandeboschia draytoniana.

wet forests at elevations of 440–1,320 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in Palmer (2003) as Sphaerocionium lanceolatum.

Hymenophyllum obtusum Hook. & Arn. palai lau li‘i ENDEMIC | SECURE

5.59 

Hymenophyllum obtusum

Small delicate epiphytes, rhizomes threadlike. Fronds 3–9 cm in length, stipes hairy, blades 2 pinnate to 3 pinnate-pinnatifid, hairs abundant on the veins and margins. Distinguished from Hymenophyllum lanceolatum by stipes and blades heavily clothed with hairs and by a shorter and more compact three-dimensional form of the fronds. Epiphytic on trees and logs in wet forest at elevations of 400–1,300 m on all of the main islands except Ni‘ihau, Kaua‘i, and Kaho‘olawe. Listed in Palmer (2003) as Sphaerocionium obtusum.

Hymenophyllum recurvum Gaudich. ‘ōhi‘a kū ENDEMIC | SECURE

Crepidomanes minutum (Blume) K. Iwats. INDIGENOUS | SECURE

5.58 

5.57 

Crepidomanes minutum

Hymenophyllum lanceolatum

Small delicate epiphytes growing in colonies on tree trunks and logs. Fronds nodding, up to 16 cm in length, arising from creeping, threadlike rhizomes; blades lanceolate, 2 to 4 pinnate-pinnatifid; stipes, rachises and costae, and veins dark brown in color; margins of pinnae and veins with simple hairs. Colony-forming epiphyte on tree trunks in

5.60 

Hymenophyllum recurvum

FERNS AND FERN ALLIES

Small delicate epiphytes found on trees and occasionally growing over mossy rocks or soil. Drooping fronds to 7–35 cm in length arising from creeping, threadlike rhizomes; blades glabrous, light green in color, 3 pinnate with the apices of both fronds and pinnae recurved. Commonly forming colonies on tree trunks in wet mossy forests at elevations of 270–1,525 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in Palmer (2003) as Mecodium recurvum.

Vandenboshia—A genus of about 20–30 species of mostly epiphytic, twining ferns found in the tropics but also in North America, southern South America, New Zealand, southern Africa, Europe, and Japan. These species would fit within a broad concept of Trichomanes. The fronds are pinnately compound, with the ultimate segments 1 veined. The indusia are cylindrical to cup shaped. The name honors Roelof Benjamin van den Bosch, a nineteenth-century Dutch botanist. Represented in the Hawaiian Islands by 3 endemic species.

Vandenboschia cyrtotheca (Hillebr.) Copel. ENDEMIC | SECURE

Vandenboschia davallioides (Gaudich.) Copel. palai hihi ENDEMIC | SECURE

Isoëtes hawaiiensis W. C. Taylor & W. H. Wagner ENDEMIC | RARE

5.62 

Vandenboschia davallioides

Medium-sized terrestrial to epiphytic ferns arising from a climbing rhizome. Fronds erect to drooping, 12–45 cm in length; blades 2 to 3 pinnate, lanceolate; stipe short, only about one-fifth the length of the frond; indusia cone shaped and stalked with a flaring tip. It differs from V. cyrtotheca by its short stipe and lanceolate blades not bent backward from the stipe. Common in shaded mesic to wet forest at elevations of 250–1,800 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Isoëtaceae

5.61 

Vandenboschia cyrtotheca

Small to medium-sized terrestrial to epiphytic ferns with climbing rhizomes. Fronds 13–37 cm in length; blades 2 to 3 pinnate, often bent back at an angle of 45 degrees from the stipe. Sori oriented at right angles to the blade. Characteristic features are long stipes that cover a third to half the length of the frond and triangular blades widest at their base. Common climbing over rocks and onto lower tree trunks in mesic to shaded wet forests at 150–750 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

lake and pond margins, while others are amphibious. Species have been shown to utilize crassulacean acid metabolism to take advantage of higher levels of dissolved CO2 at night in oligotrophic lakes. The name is from the Greek, isos, equal, and itas, year, referring to the evergreen habit. Represented in the Hawaiian Islands by 1 endemic species.

Small herbaceous perennial aquatics that are rushlike in appearance. Key features are the tufted, quill-shaped leaves that arise from a cormlike axis, the sporangium borne on the leaf side toward the axis, and the absence or rarity of any kind of vegetative reproduction. There are about 200 species, with the single genus Isoëtes cosmopolitan in distribution. A segregate genus, Stylites, is now included within Isoëtes. The family has no economic importance. In the Hawaiian Islands represented by 1 endemic species of Isoëtes.

Isoëtes—A genus of about 200 worldwide species of mostly temperate regions with poor representation in the tropics. Many species are submerged aquatics along

5.63 

Isoëtes hawaiiensis

Submerged to emergent aquatic plants with cormlike rhizomes and bearing a cluster of spirally arranged grasslike leaves. A very localized species known only from about 0.4 ha of ponds on the summit of ‘Eke on West Maui and on lava rocks of a small stream course at 850 m elevation on Mauna Kea, Hawai‘i. The ‘Eke ponds dry completely in drought years.

Lindsaeaceae

Terrestrial ferns. Key features are short to long creeping rhizomes, blades 1–3 pinnate or more divided, veins usually free. Sori marginal or submarginal indusial, typically open toward margin. There are eight genera and about 200 species,

207

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FERNS AND FERN ALLIES

widely distributed in tropical and subtropical regions. The family is sometimes included within the Dennstediaceae. Some species are cultivated as ornamentals. Represented in the Hawaiian Islands by the indigenous genera Lindsaea and Sphenomeris. A natural hybrid between Sphenomeris chinensis and Lindsaea ensifolia has been named xLindsaeosoria flynii W. H. Wagner.

Often placed into the genus Odontosoria, a group otherwise restricted to Central America, the Caribbean, and Africa. Many species form thickets with long scandent fronds that trail over vegetation. The name comes from the Greek sphen, wedge, and meros, part, referring to the wedge-shaped terminal segment of the frond. In the Hawaiian Islands, there is 1 indigenous species.

Lindsaea—A pantropical genus of about 180 tropical species of small to medium-sized ferns. Typically terrestrial but some epiphytic. Rhizomes creeping and covered with bristlely scales. Blades 1to 4-pinnate. Named in honor of John Lindsay, a nineteenth century surgeon in Jamaica. In the Hawaiian Islands represented by one indigenous species and one naturalized species.

Sphenomeris chinensis (L.) Maxon. pala‘ā, palapala‘ā, pā‘ū o Pala‘e INDIGENOUS | SECURE

Lindsaea repens (Bory) Thwaites laukahi

Nephrolepis cordifolia (L.) C. Presl. kupukupu

INDIGENOUS | SECURE

5.65 

5.64 

Lindsaea repens var. macraeana

Medium-sized terrestrial fern, occasionally epiphytic, with long creeping rhizomes. Easily recognized by its linear 1-pinnate fronds 20–50 cm in length and tapering on each end. Small discrete sori are arrayed near the blade margin and covered by small curved indusia opening outward. Uncommon in wet mossy forests at elevations of 180–765 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Hawaiian plants represent the endemic var. macraeana.

Sphenomeris—A genus of paleotropical and temperate subtropical, mostly terrestrial ferns with about 10 species.

Nephrolepis—A genus of terrestrial, rock-dwelling, or epiphytic ferns that are cosmopolitan in the tropical and warm temperate areas of the world. There are 19 species. The genus is characterized by abundant long stolons, with clustered linear fronds rising erectly. Fronds have short stipes and 1-pinnate blades, with sessile to short-stalked pinnae that are easily detached from the rachis. Indusia kidney shaped and attached at the base, opening at a 45-degree angle toward the tip of the pinnae. The Boston fern, Nephrolepis exaltata, and its cultivars are perhaps the most commonly cultivated ferns. The name is from the Greek nephros, kidney, and lepis, scale, referring to the shape of the indusia. In the Hawaiian Islands, there are 2 indigenous species and 4 naturalized species, with 2 endemic hybrids. Palmer (2003) placed this genus within the Nephrolepidaceae.

Sphenomeris chinensis

INDIGENOUS | SECURE

Medium-sized terrestrial ferns arising from a short creeping rhizome. Fronds 15–80 cm in length, blades 3 pinnate-pinnatifid to 4 pinnate; ovate-lanceolate in shape, ultimate segments wedge shaped and narrow. Sori elongate at the tips of the ultimate segments; indusia pocketlike and opening toward the margin. A very common species growing in mesic to wet forests, grasslands, and shrublands at elevations of 40–1,310 m on all of the main islands except Ni‘ihau and Kaho‘olawe. The Hawaiians used the fronds to make a red dye and interlaced the fronds with maile in lei.

Lomariopsidaceae

Terrestrial, rock-growing, and epiphytic ferns, small to moderate in size. The fronds are typically simple or once pinnate, with separate fertile and vegetative fronds. Lower blade surface of fertile fronds covered with sporangia. The family is pantropical in distribution, with 4 genera and about 70 species. The Lomariopsidaceae is sometimes included within the Dryopteridaceae. Represented in the Hawaiian Islands by 1 indigenous genus. Palmer (2003) placed Elaphoglossum within this family, but it is now included within the Dryopteridaceae.

5.66 

Nephrolepis cordifolia

Medium-sized terrestrial ferns, occasionally epiphytic. Rhizomes trailing to erect. Fronds erect to drooping, 30–70 cm in length; blades linear and uniform in width except at the base and tip; pinnae often deciduous with age. Sori medial on the pinnae, depressed on the lower

FERNS AND FERN ALLIES

surface and thus forming convex elevations on the upper surface; indusia opening toward the tip of the pinnae. Locally common in mesic to wet forests at 440– 1,525 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Nephrolepis exaltata (L.) Schott ni‘ani‘au, pāmoho, kupukupu INDIGENOUS | SECURE

differ among systematists. The family is cosmopolitan in distribution, being well represented in arctic, temperate, and tropical regions. Economically the family is of minor importance, with some species used in the florist trade or cultivated, while the spores of others were employed medicinally. Represented in the Hawaiian Islands by 3 indigenous genera: Huperzia, Lycopodium, and Lycopodiella.

Huperzia—A cosmopolitan genus of terrestrial or rock-dwelling fern allies with about 200 species that range in distribution from tropical to temperate and subarctic and alpine habitats. Characterized by clustered erect to nodding stems that are equally branched throughout but lacking a horizontal main stem or rhizome. Sporangia borne on short stalks from the axils of unmodified leaves. The genus honors Johann Peter Huperz, a nineteenth-century German horticulturist. Represented in the Hawaiian Islands by 5 endemic and 5 indigenous species, which represent 3 lineages within the genus. Hybrids between related species are relatively common.

Delicate terrestrial herbs, 4–12 cm in height, stems erect but decumbent at their base, often with 1–2 branches, pink to red in color. Leaves spreading to reflexed, narrowly lanceolate to almost linear, 2–4 mm in length. Distinguished from other species by its narrow pink stems and small lanceolate leaves. Infrequent but locally common in boggy and mossy areas of wet forest at elevations of 800–1,645 m on all of the main islands except Ni‘ihau, Lāna‘i, and Kaho‘olawe.

Huperzia phyllantha (Hook. & Arn.) Holub wāwae‘iole INDIGENOUS | SECURE

Huperzia erubescens (Brack.) Holub INDIGENOUS | SECURE

5.67 

Nephrolepis exaltata

Medium-sized terrestrial ferns arising from an erect rhizome with many stolons. Fronds erect, 30–100 cm in length; blades elliptic-lanceolate, widest in the middle and tapering to both ends, pinnae coriaceous, somewhat dimorphic with both sterile and fertile pinnae. Sori submarginal, irregularly spaced; indusia kidney shaped, opening outward at 45 degrees from the costae. Common in mesic to wet forest and high-elevation dry forest at 25–1,400 m on all of the main islands except Kaho‘olawe. The Hawaiian plants represent the endemic subsp. hawaiiensis. 5.69 

Lycopodiaceae

Low terrestrial or epiphytic fern relatives, often resembling mosses. Key features are the presence of numerous small, often spirally arranged leaves and reproduction by spores from sporangia that are borne singly on the upper side of the leaves near their base or above the leaves in a strobilus. Five genera and about 300 species are recognized, although generic concepts

5.68 

Huperzia erubescens

Huperzia phyllantha

Medium-sized epiphytes 20–70 cm tall, with stems often drooping at least in the upper part. This large size and an abrupt transition from sterile to much smaller fertile leaves separate this species from related taxa. Common epiphyte in mesic to wet forests at elevations of 250–945 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

209

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FERNS AND FERN ALLIES

Huperzia serrata (Thunb. ex Murray) Trevis. INDIGENOUS | SECURE

Lycopodiella cernua (L.) Pic. Serm. wāwae‘iole

Lycopodium venustulum Gaudich. INDIGENOUS | SECURE

INDIGENOUS | SECURE

5.71 

5.70 

Huperzia serrata

Terrestrial plants with erect stems up to 25 cm tall; foliose throughout. Sterile and fertile leaves are alike, spreading and reflexed, obovate to oblanceolate in shape, but sharply constricted at their base. The stems show a distinctive annual constriction of growth. Occurs in mesic to wet forest at elevations of 520–1,220 m on all of the main islands except Maui, Ni‘ihau, and Kaho‘olawe, but rare on Moloka‘i and Lāna‘i.

Lycopodiella—A pantropical to subtropical genus of about 40 species of terrestrial fern allies. Horizontal creeping rhizomes give rise to upright tree-form branches with both fertile and sterile leaves, with the former generally smaller. The genus is separated from Lycopodium by the presence of fertile leaves with sessile strobili nodding downward from their axils. The name is from the Greek lykos, wolf, and podes, foot, referring to Lycopodium, and -ella, a diminutive suffix. Represented in the Hawaiian Islands by a single indigenous species.

Lycopodiella cernua

Terrestrial plants with stiff, erect stems to 100 cm tall. The repeated branching, spreading or erect, terminates in a nodding strobilus of aggregated fertile leaves bearing the sporangia at their bases; leaves needlelike and spreading. Common in a variety of habitats, from mesic shrubland and grassland to wet forest and bogs, at elevations of 15–2,135 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Pantropical in distribution and probably the most abundant clubmoss in the world. Occasionally employed in the florist trade. The Native Hawaiians used a boiled preparation of this species to treat rheumatism.

Lycopodium—A genus of terrestrial plants

with 15–25 species that is cosmopolitan in distribution in mostly temperate and subarctic regions. Characterized by the presence of horizontal stems and rhizomes, with upright shoots alternating along the rhizome and fertile leaves aggregated into strobili borne on distinct stalks. The name is from the Greek lykos, wolf, and podes, foot, alluding to the resemblance of the branch tips to a wolf’s foot. Represented in the Hawaiian Islands by 1 indigenous species.

5.72 

Lycopodium venustulum

Perennial herbs with stems trailing over the ground, sparsely rooting. Stems creeping, horizontal, or upright with branches arising at about a 45-degree angle, often 1 m or more in length, with upright sterile stems clothed with needle-shaped leaves densely crowded on the branches. Upright fertile stems bearing 3–6 stobili. In wet forest and shrublands extending into the subalpine zone at elevations of 825–2,135 m. Present on all of the main islands except Ni‘ihau and Kaho‘olawe. On O‘ahu known only from near the summit of Mount Ka‘ala. There are 2 varieties with the typical form shown, and the var. verticale is endemic to Mauna Kea on Hawai‘i.

Marattiaceae

Large terrestrial ferns. Key features are the large and often leathery 1-to-many-pinnate fronds, which are commonly deciduous, and the presence of large amounts of tannin and mucilage, especially in the rootstocks. Stems and blades with scales, lenticels scattered on petioles

FERNS AND FERN ALLIES

and rachises. Sporangia free or in round or elongate synangia. A family of 4 genera and about 150 species distributed in the tropics of the Southern Hemisphere. A few species of Angiopteris and Marattia are cultivated as ornamentals. In the Hawaiian Islands represented by the indigenous genus Marattia and the introduced Angiopteris evecta, an escape from cultivation that has aggressively spread in moist valleys on O‘ahu.

Marattia—A genus of about 50 species of

large terrestrial ferns that are pantropical and subtropical in distribution. These ferns have large, starchy rootstocks and often very large 1- or 4-pinnate fronds. The name honors eighteenth-century botanist G. F. Maratti. In the Hawaiian Islands, there is 1 endemic species.

Marratia douglasii (C. Presl) Baker pala, kapua ‘ilio ENDEMIC | SECURE

but now restricted to areas without feral pigs. The thick, fleshy rootstocks are rich in starch and mucilage. Formerly used by the Hawaiians as a food source and as a remedy for bronchitis and diarrhea.

Marsileaceae

(CLOVER FERN FAMILY)

Aquatic or partially submerged wetland ferns. Key features are the distinctive fronds with 4, 2, or no leaflets at the apex of the petiole and the sori borne within sporocarps near the base of the plant. The family is cosmopolitan in distribution, with 3 genera and 75 species (70 of these in Marsilea). Some species are occasionally cultivated in ponds or aquaria. Represented in the Hawaiian Islands by the indigenous genus Marsilea.

Marsilea—A cosmopolitan genus of ferns

in marsh or aquatic habitats with about 70 species. The fronds consist of 2 adjacent pairs of leaflets attached at the tip of the petiole; the sori are enclosed in a hardened sporocarp attached near or at the base of the petiole. The name honors Count Luigi Ferdinando Marsigli, a seventeenth-century Italian botanist. In the Hawaiian Islands, there is 1 endemic species.

Marsilea villosa Kaulf. ‘ihi‘ihi, ‘ihi lā‘au, water clover fern ENDEMIC | ENDANGERED

5.73 

Marattia douglasii

Medium-sized to large terrestrial ferns. Rhizomes form tall stems up to 55 cm tall; covered with large, fleshy stipules at the base of the stipes. Fronds 90–270 cm in length; blades 3 to 4 pinnate. The clusters of fused sori termed synangia are borne along the vein margins on the lower blade surface. Only the aggressive alien Angiopteris evecta shares these features, but it has much larger fronds. Once common in mesic to wet forests at elevations of 425–2,070 m on all of the main islands except Ni‘ihau and Kaho‘olawe

borne in a separate fruiting body, called a sporocarp, near the rhizome base. A rare plant known today from aquatic habitats from 5–6 low-elevation sites on O‘ahu and several sites on Moloka‘i. Extinct on Ni‘ihau.

Nephrolepidaceae—This family, consisting of the single genus Nephrolepis, is now included within the Lomariopsidaceae.

Ophioglossaceae

(GRAPEFERN AND ADDER TONGUE FAMILY)

Terrestrial ferns, occasionally epiphytic. Key features are the presence of a solitary leaf or a few leaves that do not exhibit circinnate vernation. They are dimorphic, usually with a sterile leafy segment and a fertile spore-bearing branch arising from a common stem. The family is cosmopolitan in distribution and is composed of 4 genera and about 80 species. Some older references term this family as the Botrychiaceae. Species of Ophioglossum are occasionally cultivated. Represented in the Hawaiian Islands by the indigenous genera Botrychium and Ophioglossum.

Botrychium—A genus of 50–60 species that are cosmopolitan in distribution but with the greatest diversity in high latitudes and at high elevations in meadows and open woods. Stems are erect, with blades 1–5 pinnate. The name comes from the Greek botrys, grape, which is the basis for the common name “grapeferns” in reference to the sporangia clusters. Represented in the Hawaiian Islands by the endemic Botrychium subbifoliatum, which was likely always rare and is thought to be extinct today. Listed in Palmer (2003) as Sceptridium subbifoliatum. Ophioderma—This genus is now considered to be part of Ophioglossum.

Ophioglossum—A genus of 25–30 species

5.74 

Marsilea villosa

Terrestrial aquatic ferns arising from a rhizome covered with reddish hairs and fronds composed of 4 leaflets resembling a four-leaf clover. The sporangia are

of ferns that are cosmopolitan in distribution. In these ferns, the sterile and fertile portions of the frond are entire. The name is from the Greek ophis, snake, and glossa, tongue, referring to the fertile portion of the frond. These are commonly called adder tongue ferns. In the Hawaiian Islands represented by 4 indigenous species.

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FERNS AND FERN ALLIES

Ophioglossum pendulum L. puapua moa INDIGENOUS | SECURE

on open sand dunes, loose lava rocks, and grasslands from sea level to 160 m on all of the main islands except Ni‘ihau. Colonies of up to hundreds of individuals are known to appear in January and February after heavy rains. Reported from old house and midden sites of early Hawaiian occupation.

Polypodiaceae

(LICORICE FERN FAMILY)

5.75 

Ophioglossum pendulum

Medium-sized terrestrial ferns arising from an inconspicuous short and tuberous rhizome. Fronds entire, strap- or ribbonlike, pendent, lacking a midrib, 20–180 cm in length. Fertile spikes pendent, generally arising from the basal half of the fronds. Common epiphyte on trees in mesic to wet forests at elevations of 40–1,000 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in Palmer (2003) as Ophioderma pendulum.

Ophiglossum polyphyllum A. Braun pololei, adder’s tongue INDIGENOUS | SECURE

Terrestrial and epiphytic ferns. Key features are blades that are mostly simple to pinnatifid or 1 pinnate and round sori with the sporangia borne in long lines or distributed along the veins. There is no indusium. Some genera have dimorphic fronds. A diverse and cosmopolitan family with 56 genera and over 1,200 species, the majority of which are epiphytes. Many species are grown as ornamentals. In the Hawaiian Islands represented by the endemic genus Adenophorus and the indigenous genera Grammitis, Lepisorus, Microsorum, Polypodium, and Stenogrammitis. Phlebodium, Phymatosorus, and Platycerium are introduced and naturalized. The Grammitidaceae was considered to be a separate family by Palmer (2003) but is now placed within the Polypodiaceae.

Adenophorus—An endemic genus of small

Small and delicate epiphytic ferns with a short creeping rhizome. This is the smallest and most delicate species in the genus. Fronds erect and clustered, 4–8 cm in length, blades 2 pinnate, ultimate segments radiating from the plane of the blade in living plants. Sori wider than ultimate segments. Infrequent to locally common in wet forests at elevations of 510–1,310 m on Kaua‘i, O‘ahu (Ko‘olau Mountains), Lāna‘i, and Maui.

Adenophorus epigaeus (L. E. Bishop) W. H. Wagner ENDEMIC | SECURE

Small and delicate epiphyte growing over mosses at the base of trees or occasionally epiphytic on low branches. Rhizomes long creeping and slender, 1.0–1.5 mm in diameter. Fronds erect, well separated, 8–18 cm in length, Blades 2 pinnate, linear to sublinear. Sori wider than ultimate segments. Similar to A. tamariscinus but distinguished by its rhizome morphology and well-separated fronds. Locally common in wet forests and bogs at elevations of 1,050– 1,550 m on Kaua‘i (see figure 2.6).

Adenophorus hymenophylloides (Kaulf.) Hook. & Grev. pai, palai huna ENDEMIC | SECURE

to medium-sized epiphytic ferns consisting of 10 species with 2 interspecific hybrids. The fronds are simple to 3 pinnatifid. Two distinct subgenera are present, with one forming clones by rhizome growth and branching and the other by root proliferations. Hybrids exist between species in each subgenus but not between subgenera. The name comes from the Greek adenos, gland, and phoros, bearing, referring to the presence of the glandular hairs.

Adenophorus abietinus (D. C. Eaton) K. A. Wilson ENDEMIC | SECURE

5.76 

Ophioglossum polyphyllum

Small terrestrial ferns arising from a subterranean rhizome. Fronds dimorphic; sterile blades erect, usually 1–2, elliptic to lanceolate in shape, 2–8 cm in length. Fertile blades 1–2 times the length of the sterile blades, with 12–20 pairs of sporangia at their tips. Scattered in occurrence but may be locally and seasonally common

5.77 

Adenophorus abietinus

5.78 

Adenophorus hymenophylloides

FERNS AND FERN ALLIES

Small and delicate epiphytic ferns. Vegetative reproduction by branching rhizomes. Fronds erect and clustered, 4–8 cm or more in length; blades 2 pinnate, ovate to lanceolate in shape, ultimate segments radiating at an angle from the plane of the blade. Locally common on trees in wet forest at elevations of 350– 1,750 m on all of the main islands except Ni‘ihau and Kaho‘olawe. This species is known to produce hybrids with A. tamariscinus and A. tripinnatifidus.

Adenophorus pinnatifidus Gaudich.

Adenophorus periens L. E. Bishop palai lā‘au

Small epiphytic ferns arising from a short rhizome. Fronds erect, arching, or pendulous, 5–12 cm in length; blades dark green, widest at their middle, deeply pinnatifid, margins glabrous. Locally common species, usually growing on tree trunks but occasionally on mossy rocks, in mesic and wet forests at elevations of 240–1,680 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Two varieties are recognized, with the widespread var. pinnatifidus shown. The var. rockii is known from Kaua‘i, the Ko‘olau Mountains of O‘ahu, and from Moloka‘i and Maui.

ENDEMIC | ENDANGERED

ENDEMIC | SECURE

Adenophorus tenellus (Kaulf.) Ranker kolokolo ENDEMIC | SECURE

5.80 

Adenophorus pinnatifidus var. pinnatifidus

Adenophorus tamariscinus (Kaulf.) Hook. & Grev. wahine noho mauna ENDEMIC | SECURE

5.82 

Adenophorus tenellus

Small epiphytic ferns arising from a slender creeping rhizome. Often forming tangled rhizome mats. Fronds simple, linear, 3–20 cm in length; blades narrow and long linear, pale green, and chartaceous in texture. Sori sparse and scattered, often extending beyond the blade margin. A common species forming epiphytic mats in mesic to wet forest at elevations of 365– 1,675 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in Palmer (2003) as Grammitis tenella.

Adenophorus tripinnatifidus Gaudich. 5.79 

ENDEMIC | SECURE

Adenophorus periens

Small epiphytic ferns arising from a short, trailing rhizome. Fronds erect to arching, 10–40 cm in length; blades pendulous, linear in shape and uniform in width, deeply pinnatifid, cut almost to the costae. Vegetative reproduction by root proliferations. Once widely occurring as an epiphyte in wet forest at elevations of 700–1,300 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Today known from only a few locations on Kaua‘i, Moloka‘i, and Hawai‘i. Currently known from 11 populations with estimates ranging from 200 to 2,000 extant plants. Most of these are in a single population at Kahauale‘a, Hawai‘i. Last recorded on O‘ahu in the early 1900s. Threats come from dieback of host trees, feral pigs, and alien plant species.

5.81 

Adenophorus tamariscinus var. tamariscinus

Small and delicate epiphytic ferns arising from a short creeping and thick rhizome covered with dark brown shining scales. Fronds clustered and erect, typically 8–20 cm in length; blades dark green, bipinnate, often with pinnae radiating away from the plane of the blade. Vegetative reproduction by branching rhizomes. Common epiphyte, especially on ‘ōhi‘a in mesic and wet forest at elevations of 300– 1,300 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Two varieties are recognized, with the widespread var. tamariscinus shown. The var. montanus is restricted to Moloka‘i, Maui, and the Kohala Mountains on Hawai‘i.

5.83 

Adenophorus tripinnatifidus

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FERNS AND FERN ALLIES

Small epiphytic ferns, neither terrestrial nor epiphytic but growing over mats of moss at the base of trees. Rhizomes creeping, slender, and long. Fronds 12–25 cm in length; blades 2 to 3 pinnate, ovate-lanceolate to deltoid in shape, with small reddish to reddish brown glands. Occasional to locally common growing over moss mats in wet forest at 600–1,950 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Grammitis—A large pantropical and austral genus with about 400 species, but estimates vary depending upon authors. Several subgenera included in this genus are recognized at the generic level. These ferns are mostly small, usually epiphytic, with the fronds entire to 3 pinnate and often bearing trichomes. Sori generally arranged in single rows on each side of the midrib but scattered in some species. The name is from the Greek gramme, line, referring to the elongate sori in some of the species. This genus was formerly placed in the Grammitidaceae, a family that has now been included within the Polypodiaceae. In the Hawaiian Islands, there are 2 endemic and 1 indigenous species.

Lellingeria—Lellingeria saffordii, listed in Palmer (2003) as an endemic Hawaiian species, has now been moved to the new genus Stenogrammitis.

Lepisorus—A genus of about 40 species of epiphytic, rock-dwelling or terrestrial ferns distributed across the paleotropics to eastern Russia and the Hawaiian Islands. The sporangia are arranged in 1 row on each side of the midvein and covered by scales when young. The name is from the Greek lepis, scale, and soros, heap, referring to the scales that cover the young sori. In the Hawaiian Islands represented by 1 indigenous species.

Microsorum—A genus of about 70 species of epiphytic, rock-dwelling, or terrestrial ferns of the paleotropics and adjacent warm temperate areas. The blades are simple, pinnate, or appearing palmately lobed. The sori are scattered irregularly over the entire undersurface of the blade. The name is from the Greek micros, small, and soros, heap, referring to the small sporangial aggregations. In the Hawaiian Islands, there is 1 endemic species.

Microsorum spectrum (Kaulf.) Copel. pe‘ahi ENDEMIC | SECURE

Lepisorus thunbergianus (Kaulf.) Ching pākahakaha, ‘ēkaha ‘ākōlea INDIGENOUS | SECURE

5.86 

Microsorum spectrum var. pentadactylum

5.87 

Microsorum spectrum var. spectrum

Grammitis hookeri (Brack.) Copel. maku‘e lau li‘i INDIGENOUS | SECURE

5.85 

5.84 

Grammitis hookeri

Small epiphytic ferns arising from a thick erect rhizome. Fronds 5–20 cm in length; blades broadly linear, widest in the middle, covered with hairs. Sori in lines close to the costae. Typically growing on low mossy branches but sometimes on mossy rocks in mesic to wet forest at elevations of 800–1,750 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Also in Australia and the Malesian region.

Lepisorus thunbergianus

Small epiphytic, rock-dwelling, or terrestrial ferns arising from a slender creeping rhizome. Fronds in clusters, 6–35 cm in length; blades simple, linear-lanceolate in shape, coriaceous, glabrous, margins entire; stipes short. The sori round to oblong and covered by many umbrella-shaped glandular structures called paraphyses. A common epiphyte on trees and rocks in a range of dry to wet forests at elevations of 10–2,100 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Also occurs in Africa and Asia.

Terrestrial ferns arising from a vinelike creeping and branched rhizome. Fronds 10–50 cm in length; blades thin, 3–5 lobed to form a triangle-shaped blade, veins prominent, joining together to form areoles. Sori round and small, scattered on veins in the areoles. Infrequent terrestrial to epiphytic species of mesic to wet forests at 180–1,050 m elevation in wet forest on all of the main islands except Ni‘ihau and Kaho‘olawe. Often present as a terrestrial ground cover, with portions of the rhizomes climbing low onto tree trunks, where the fronds become epiphytic. Two varieties are present: var. pentadactylum is restricted to Kaua‘i and East Maui, and var. spectrum is widespread.

FERNS AND FERN ALLIES

Polypodium—A genus of terrestrial, rock-dwelling, or epiphytic ferns that are cosmopolitan in distribution, especially in the tropics and subtropics. There are about 110 species. The fronds disarticulate, leaving a scar above the base of the petiole. The sori are borne at the tip of a veinlet or at the junction of veins. The name is from the Greek polus, many, and podus, feet, referring to the knob-shaped prominences on the rhizome. In the Hawaiian Islands, there is 1 endemic species.

Polypodium pellucidum Kaulf. ‘ae, ‘ae lau nui, polypody fern ENDEMIC | SECURE

to 55 cm in length; blades deeply pinnatifid with pinnae divided to near the rachis. When growing in open situations, the segments of the frond may be twisted in a plane other than that of the axis. Highly variable in habit and morphology. Occurs in diverse habitats from mesic to wet forest to subalpine and alpine forest at elevations from 150–2,135 m on all of the main islands except Ni‘ihau and Kaho‘olawe Three varieties are present, with the widespread var. pellucidum and var. volcanicum (Moloka‘i, Maui, and Hawai‘i) shown. The var. acuminatum is restricted to Kaua‘i (Waimea and Hanalei) and East Maui (Kaupō Gap).

Stenogrammitis—A genus of 24 species of epiphytic and epilithic ferns with an erect to short creeping rhizome. Pantropical in distribution with species in the neotropics, Africa, Madagascar, and the Pacific Islands. Characteristic of montane cloud forests. Distinguished from Lellingeria, where these species were formerly placed, by linear leaves with unbranched veins in the segments and iridescent rhizome scales. Represented in the Hawaiian Islands by a single endemic species.

Stenogrammitis saffordii (Maxon) Labiak kihe ENDEMIC | SECURE

halves of the blades have entire margins and are covered by sori on their lower surface. Common in cloudy wet forests and bogs at elevations of 900–1,800 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed as Lellingeria saffordii by Palmer (2003).

Psilotaceae

(WHISK FERN FAMILY)

Epiphytic or terrestrial herbaceous fern allies. Key features are the dichotomous branching, a lack of leaves, the presence of slender rhizomes, and sporangia fused into a 3-lobed synangia. There are 2 genera, Tmesipteris and Psilotum, with 3 to about 15 species depending on different authors.

Psilotum—A genus of 2 species of erect or pendent mostly epiphytic perennials found throughout the tropics and subtropics of the world. Characteristics same as for the family. The name comes from the Greek psilos, naked or smooth, in reference to the smooth aerial stem. In the Hawaiian Islands, there are 2 indigenous species and an endemic hybrid. These species are easily separated by the shape of the stems, being either flattened or triangular in cross section.

Psilotum complanatum Sw. moa, moa nahele INDIGENOUS | SECURE

5.88 

Polypodium pellucidum var. pellucidum

5.89 

Polypodium pellucidum var. volcanicum

Medium-sized terrestrial ferns arising from a long creeping rhizome. Sometimes present in colonies of fronds, with rhizomes climbing lower tree trunks to become epiphytes. Fronds coriaceous, up

5.90 

Stenogrammitis saffordii

Small epiphytic ferns. Fronds simple with linear blades characterized by a prominently lobed margin on their lower half; blades 2.5–12.5 cm in length. The upper

5.91 

Psilotum complanatum

215

216

FERNS AND FERN ALLIES

Pantropical epiphyte with dichotomous branching and flattened stems that are usually pendulous on the trunks of trees. Locally common in mesic to wet forest at elevations of 250–1,100 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Psilotum nudum (L.) P. Beauv. moa, moa nahele, pipi INDIGENOUS | SECURE

were once placed in their own families, as with Ceratopteris (water fern) and the Vittariaceae (shoestring ferns). The family has been split into a number of tribes, not always with a consensus of agreement among systematists. Many genera, such as Pteris, Adiantum, Pityrogramma, and others, are often cultivated as ornamentals. In the Hawaiian Islands represented by the indigenous genera Adiantum, Coniogramme, Doryopteris, Haplopteris, Pellaea, and Pteris. Naturalized genera include Ceratopteris, Cheilanthes, and Pityrogramma.

Adiantum—A cosmopolitan genus of terrestrial or rock-loving ferns comprising about 200 species throughout the tropics and temperate regions of the world. The genus includes a remarkable leaf morphology. The sori are borne on fertile veins that enter the indusium. Many species are cultivated as ornamentals. The name is from the Greek a, without, and diane, unwetted, referring to the ability of the pinnae to shed raindrops. The Hawaiian Islands have 1 indigenous species and 4 naturalized species.

and simply pinnate above, with fan-shaped pinnae; veins end in marginal teeth. Sori are elongate in shape and borne at lobe tips of the pinnae. Uncommon but found on shaded embankments or in wet ravines from sea level to 430 m elevation on all of the main islands except Kaho‘olawe. The dark stipes were once woven in lau hala mats to create designs.

Coniogramme—A genus of paleotropical

terrestrial ferns with large once-pinnate to 3 pinnate fronds with entire to finely toothed pinnae. The elongate sori occur along the veins, except near the margin, and lack an indusium. There are about 40 species that occur from Africa east to Polynesia and Japan. The name is from the Greek konios, dust, and gramme, line, referring to the arrangement of the sporangia along the veins. In the Hawaiian Islands, there is 1 endemic species.

Coniogramme pilosa (Brack.) Hieron. lo‘ulu ENDEMIC | SECURE

Adiantum capillus-veneris L. ‘iwa‘iwa, maidenhair fern, venus’ hair fern INDIGENOUS | SECURE

5.92 

Psilotum nudum

Pantropical terrestrial to occasional epiphyte with successively dichotomously forked branches that are triangular in outline, at least below. Occurs in a wide variety of habitats from sea level to 1,200 m elevation on Midway and all of the main islands, often becoming weedy in urban gardens.

5.94 

Medium-sized terrestrial ferns with creeping rhizomes. Fronds 30–90 cm in length with a long stipe; a distinctive light green color of the blade. Blades once pinnate except for basal 1–2 pinnae with large pinnules. Long linear sori arrayed along all of the veins of fertile fronds. Locally common in mesic to wet forest at elevations of 610–2,070 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Pteridaceae

Terrestrial ferns, occasionally epiphytic or growing over rocks. Key features are fronds that are typically multipinnate but entire in the vittarioids. Linear sori on the margins of the blades. The sori lack a true indusium and instead are protected by a false indusium formed from the curved margin of the leaf. The family has a cosmopolitan distribution with about 50 genera and 950 species almost worldwide in distribution but most abundant and diverse in tropical and subtropical arid regions. Several genera included here

Coniogramme pilosa

5.93 

Adiantum capillus-veneris

Small and delicate terrestrial or rock-­ dwelling ferns with creeping rootstocks covered with brownish scales, stipes black. Fronds erect or spreading, 15–50 cm in length, blades 1 to 3 pinnate at the base

Doryopteris—A cosmopolitan genus of tropical terrestrial ferns with 40–50 species found mostly in moist or seasonally dry rocky places. These ferns are often variable in leaf shape, with relatively simple juvenile leaves grading into multilobed mature fertile fronds. The name is

FERNS AND FERN ALLIES

derived from the Greek doryos, halberd shaped, and pteris, fern, referring to the shape of the fronds of some species. In the Hawaiian Islands, there 4 endemic species.

Doryopteris decipiens (Hook.) J. Sm. kumuniu, ‘iwa‘iwa, manawahua ENDEMIC | SECURE

Small to medium-sized terrestrial ferns arising from short rhizomes. Fronds up to 30 cm in length; blades triangular in shape, pinnate to bipinnate; pinnae spreading out from the plane of the blade to give a three-dimensional appearance in living plants, ultimate segments with parallel sides; rachises winged between the 2 lowest pinnae. Occurs in dry shrublands, grasslands, and forest at elevations of 120–700 m on all of the main islands except Ni‘ihau. Often growing on exposed basalt.

fern, referring to the simple strap-shaped fronds of the genus. Represented in the Hawaiian Islands by a single indigenous species.

Haplopteris elongata (Sw.) E. H. Crane ‘ohe‘ohe, mana INDIGENOUS | SECURE

Doryopteris takeuchii (W. H. Wagner) W. H. Wagner ENDEMIC | RARE

5.95 

Doryopteris decipiens

Small to medium-sized terrestrial ferns arising from a thick rhizome. Fronds up to 30 cm in length; blades triangular in shape and flattened in a single plane, 1 pinnate-pinnatisect to 2 pinnate; rachises winged between the lowest pairs of pinnae; upper segments triangular and tapering to a rounded tip. Sori along the outer edge of the segments. Occurs in dry shrublands, grasslands, and forest at elevations of 30–915 m on all of the main islands, as well as Lehua. Often growing out of exposed basalt.

Doryopteris decora Brack. ENDEMIC | SECURE

5.98 

5.97 

Doryopteris takeuchii

Small terrestrial xerophytic ferns arising from a thick rhizome bearing a few fronds 10–30 cm in length among the older dried ones; rachises between the 2 lowest pinnae lack wings. Sori found along the outer edge of the segments. Found only on Diamond Head Crater on O‘ahu on dry rocky outcrops at elevations of 110–120 m.

Haplopteris—A genus of 5 species dis-

5.96 

Doryopteris decora

tributed from Africa across Southeast Asia and Australia to the Pacific. Small to medium-sized epiphytes with simple linear blades. Members of this genus were formerly placed within Vittaria in the Vittariaceae. The name comes from the Greek haplos, simple, and pteris,

Haplopteris elongata

Small epiphytic ferns, occasionally growing over rocks. Fronds strap shaped and linear with entire margins, virtually lacking a stipe and reaching up to 50 cm in length; clustered fronds hang pendulously. Long, linear sori present in marginal grooves on the lower blade surface. Occurs in mesic to wet forests at elevations of 20–900 m on all of the main islands except Ni‘ihau and Kaho‘olawe. At the drier end of its range, it often grows from the rhizome base of Asplenium nidus, giving the latter the appearance of having aerial roots.

Pellaea—A cosmopolitan genus of typically terrestrial or rock-dwelling ferns with about 50 species that are usually found in xeric habitats. The fronds are often leathery, 1–3 pinnate, and have a continuous marginal indusium covering several to many sori. A few species are cultivated. The name is from the Greek pellos, dusky, referring to the color of the stipes. In the Hawaiian Islands, there is 1 indigenous species.

217

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FERNS AND FERN ALLIES

Pellaea ternifolia (Cav.) Link kalamoho lau li‘i, laukahi

Pteris cretica L. ōali

INDIGENOUS | SECURE

INDIGENOUS | SECURE

Large terrestrial ferns with a short creeping rhizome. Fronds 100–250 cm in length; blades once pinnate-pinnatisect; triangular in shape; pinnae large, 20–50 cm in length. Common in local areas of wet forests at 300–2,075 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Pteris hillebrandii Copel. ENDEMIC | SECURE

5.100 

Pteris cretica

Medium-sized terrestrial ferns with short creeping rhizomes. Fronds 17–90 cm in length; blades once pinnate with the basal pinnae forked into 2 subequal branches, ovate to ovate-oblong in shape, light green in color, chartaceous, and glabrous. Stipes half or more of frond length. Locally common in dry to mesic areas at 340–2,450 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Pteris excelsa Gaudich. waimakanui, ‘iwa INDIGENOUS | SECURE

5.99 

Pellaea ternifolia

Pteris hillebrandii

Medium-sized terrestrial ferns. Fronds to 85 cm in length, with the stipe comprising half or more of this length. Blades once pinnate distally and basally with 2–3 pinnae unequally forked into branches. Rachises winged in upper portions. Resembles P. cretica but blades variably and irregularly dissected. Uncommon in mesic forest at elevations of 370–1,220 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Small terrestrial or rock-dwelling xerophytic ferns arising from a thick, branching rootstock; fronds tufted, 9–40 cm in length, with shiny dark stipes. The paired pinnae are coriaceous, divided to the base into 3 linear-elliptic segments. The uppermost pairs sometimes 2 cleft and the terminal one usually simple. Found at elevations of 600–3,500 m on all of the main islands except Ni‘ihau and Kaho‘olawe but rare outside of subalpine and alpine areas of Maui and Hawai‘i. Also native to North and Central America.

Pteris—A large cosmopolitan genus comprising about 250 species of terrestrial ferns, with a rhizome that is short creeping or erect. Stipe strawlike, reddish, brown or purple, scaly at the base. Blades 1–4 pinnate or basally divided into 3 parts. Marginal to submarginal linear sori connecting the vein endings. The name comes from the Greek pteris, fern, an ancient name referring to ferns in general. In the Hawaiian Islands represented by 3 endemic and 2 indigenous species.

5.102 

Pteris irregularis Kaulf. mānā, ‘āhewa, ‘iwa puakea ENDEMIC | SECURE

5.101 

Pteris excelsa

5.103 

Pteris irregularis

FERNS AND FERN ALLIES

Medium-sized to large terrestrial ferns with a short caudex. Fronds 60–100 cm or more in length; stipes half or more of frond length; blades once pinnate-pinnatifid to 2 pinnate-pinnatifid, ovate to ovate-triangular in shape, chartaceous, pinnae cut to winged rachises. Locally common in dry to mesic forest at elevations of 450–1,920 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Schizaeaceae

Terrestrial ferns, often small and grasslike or sometimes climbing. Key features are sporangia not in discrete sori and borne on marginal branched or unbranched projections at blade tips; indusial lacking. Blades linear or fan shaped. There are 2 genera and about 30 species, with distribution centered in the tropics and warm temperate Southern Hemisphere. Lygodium is often cultivated for its climbing habit. In the Hawaiian Islands represented by the indigenous genus Schizaea and the introduced and now naturalized genus Lygodium.

Schizaea—A genus of 28 species of mostly tropical terrestrial ferns. The fronds are small and dimorphic with tufted brushor comb-shaped tips. Fertile fronds have small blades at their tips and sporangia arrayed under the revolute margins of the pinnae. The name is from the Greek schizos, deeply divided, referring to the deeply cleft fronds of some species. In the Hawaiian Islands, there is 1 endemic species.

Schizaea robusta Baker ‘oāli‘i makali‘i, haili o Pua ENDEMIC | SECURE

Small terrestrial ferns with wiry stipes up to 30 cm long. Fronds closely clustered, variable in length from 3–42 cm, sometimes resembling a toothbrush, with a long stipe and the pinnae all turned to one side. Dense numbers of stipes lacking blades at the rhizome tip give the appearance of a wiry mat. An uncommon species in bogs or sometimes open areas in wet forest at elevations of 500–1,750 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Medium-sized plants with erect stems that branch from the middle into a dense pyramidal frond flattened in one plane. Strobili bearing the sporangia are small and confined to the tips of the outer branchlets of the fronds, which may reach 20–60 cm in length. Common on shaded rocks, soil, and moist cliff faces at elevations of 100–1,200 m in mesic to wet forest on all of the main islands except Ni‘ihau and Kaho‘olawe. A variable plant throughout its range.

Selaginellaceae

Selaginella deflexa Brack.

Mosslike or clubmosslike perennial fern allies. Key features are the stems with many dichotomous branches, the arrangement of the leaves in spirals (opposite pairs or rows of 4), the production of 2 kinds of spores, and the presence of stobili in all species. The single genus Selaginella, with about 700 species, is cosmopolitan in distribution, including the Hawaiian Islands. Some species are grown as ornamentals, especially the peacock plant, S. wildenovii.

ENDEMIC | SECURE

Selaginella—A genus of about 700 species

of clubmosslike plants that are nearly worldwide in distribution. Sometimes they can superficially resemble some of the clubmosses, mosses, or liverworts. Plants terrestrial or growing on rock; only rarely epiphytic. Separated from the true clubmosses by differences in the sporophyte and gametophyte and by the presence of a strap-shaped ligule at the base of the leaves. The name is from the diminutive selago, a classical name for some species of Lycopodium. In the Hawaiian Islands, there are 2 endemic species and 3 naturalized species.

Selaginella arbuscula (Kaulf.) Spring lepelepe a moa ENDEMIC | SECURE

5.106 

Selaginella deflexa

Small plants with slender stems dividing at the base into several erect branches that are leafy throughout. The lower sterile leaves are green, while the upper fertile ones are straw colored. Distinct from other Hawaiian species of Selaginella because of its small size, needle-shaped leaves with pointed marginal lobes, and strobili that are wider than the sterile stems. Infrequent but locally common in wet swampy places and open mossy bogs at elevations of 1,050–1,500 m on all of the main islands except Lāna‘i, Ni‘ihau, and Kaho‘olawe.

Tectariaceae

5.104 

Schizaea robusta

5.105 

Selaginella arbuscula

Terrestrial ferns. Rhizomes covered with lanceolate scales; usually erect but rarely prostrate. Blades typically simple, pinnate, or bipinnate. Sori along the veins, and indusium kidney shaped or peltate,

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occasionally absent. Pantropical in distribution with 8–15 genera—but generic limits are uncertain—and about 400 species. Represented in the Hawaiian Islands by the indigenous genus Tectaria.

Tectaria—A genus of terrestrial or rock-­ dwelling ferns that are found throughout the tropics and subtropics of the world. There are about 150 species. The fronds are usually all of one kind, sometimes dimorphic. The blades may be simple to lobed and 1 to 3 pinnate. The sori are usually found along the veins. The name is from the Latin tectus, covered or hidden, referring to the protecting indusium. The Hawaiian Islands have 1 endemic species and 1 naturalized species.

Tectaria gaudichaudii (Mett.) Maxon ‘iwa‘iwa lau nui ENDEMIC | SECURE

Thelypteridaceae

Terrestrial ferns. Key features include the presence of 2 ribbon-shaped vascular bundles visible in a cross section at the base of the stipe. The midrib and blades of the fronds have short, single-celled, sharp-tipped hairs on their upper surface. Sori round to oblong in shape, rarely elongate, along veins, with kidney-shaped indusia or occasionally lacking indusia. Almost cosmopolitan in distribution. Some authors recognize 1 genus, while others see 5–30 genera. An intermediate view of the family includes 6 genera and about 950 species. Some species are grown as ornamentals. Represented in the Hawaiian Islands by 3 indigenous genera: Cyclosorus, Pseudophegopteris, and Thelypteris. Palmer (2003) divided the Hawaiian members of the family differently into 5 genera. Other authors have lumped all of these within a broad concept of Thelypteris. The naturalized genus Macrothelypteris also occurs in the Hawaiian Islands.

Medium-sized to large terrestrial ferns arising from a thick rhizome to 3 cm in diameter. Fronds variable in size but may reach up to 2 m in length, with linear, once-pinnate blades with a toothed margin. A common fern of diverse habitats from forest floors to shaded stream banks, steep rock walls, and waterfalls at elevations of 120–1,400 m on all of the main islands except Ni‘ihau and Kaho‘olawe. The Hawaiians used the plants both medicinally and as a vegetable cooked with meat and kalo. Listed by Palmer (2003) as Christella cyantheoides.

Cyclosorus interruptus (Willd.) H. Itō INDIGENOUS | SECURE

Amauropelta—See Thelypteris. Christella—See Cyclosorus. Cyclosorus—A genus of about 600 species

of tropical terrestrial ferns. The name comes from the Greek kyklos, circle, and soros, heap, referring to the round sori of the genus. In the Hawaiian Islands represented by 5 endemic and 1 indigenous species, with 2 additional species naturalized. Palmer (2003) treats these species as members of the genera Christella and Pneumatopteris.

Cyclosorus cyatheoides (Kaulf.) Farw. kikawaiō ENDEMIC | SECURE

5.109  5.107 

Tectaria gaudichaudii

Medium-sized to large terrestrial ferns arising from a thick prostrate rhizome. Fronds 30–140 cm in length, clustered; blades once pinnate-pinnatifid to bipinnate; triangular in shape, dark green, membranous; stipes glossy dark brown, 20–70 cm in length. Stipes may be 2–7 cm long. Occurs in shaded places in valleys and gorges in mesic and wet forest at 210–1,250 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

5.108 

Cyclosorus cyantheoides

Cyclosorus interruptus

Medium-sized terrestrial fern with creeping subterranean rhizome heavily covered in branching, hairy roots. Fronds 45–60 cm in length, stipes one-third to half frond length. Blades once pinnate, pinnae with small red glands on their lower surface, lower surface of costae with broad triangular scales. Sori forming almost continuous submarginal lines, indusia kidney shaped. Occurs in freshwater marshes, bogs, and abandoned taro patches at elevations of sea level to

FERNS AND FERN ALLIES

1,070 m on all of the major islands except Ni‘ihau and Kaho‘olawe.

Cyclosorus sandwicensis (Brack.) Copel. hō‘i‘o kula

Pseudophegopteris keraudreniana (Gaudich.) Holttum waimakanui, ‘ākolea, ala‘alai

Thelypteris globulifera (Brack.) C. F. Reed palapalai a Kamapua‘a ENDEMIC | SECURE

ENDEMIC | SECURE

ENDEMIC | SECURE

5.112 

5.110 

Cyclosorus sandwicensis

5.111 

Pseudophegopteris keraudreniana

Pneumatopteris—See Cyclosorus.

Terrestrial, scandent ferns arising from an underground rhizome. Fronds originate 4–8 cm apart along the rhizome and exhibit indeterminant viny growth up to 6 m or more in length; blades 2 pinnate-pinnatifid to 2 pinnate-pinnatisect; characterized by 2 crescent-shaped vascular bundles in a cross section of the stipe. Locally common in wet forests at elevations of 200–1,800 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Pseudophegopteris—A genus of large terres-

Thelypteris—In its broad interpretation,

Medium-sized terrestrial ferns arising from a stout, erect rhizome. Fronds 20–100 cm in length, blades once pinnate, lanceolate, rachis covered with white multicellular hairs. Common in mesic to wet forests, especially on stream banks, at elevations of 750–2,100 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed in Palmer as Pneumatopteris sandwicensis.

trial ferns with 20 species found from the island of St. Helena off the coast of Africa east to Polynesia and Southeast Asia. The bipinnate fronds have widely spaced pinnae along the stipe. The sori are without an indusium and often spread along the veins. The name is from the Greek pseudo, false, and Phegopteris, a genus of north temperate ferns. In the Hawaiian Islands, there is 1 endemic species.

a very large and complex genus of typically terrestrial ferns that includes the entire family, but it is more commonly now split into multiple genera. Sori round, oblong, or elongate along veins. The name comes from the Greek thelys, female, and pteris, fern, referring to a delicate fern. As interpreted here, represented in the Hawaiian Islands by a single endemic species.

Thelypteris globulifera

Medium-sized terrestrial ferns. Rhizomes trailing to erect, covering old stipes. Fronds 40–140 cm in length; stipes short and covered with white hairs; blades once pinnate-pinnatifid, oblanceolate, with glands on the under surface, gradually tapering to small pinnae at both ends. It is similar in appearance to several species of Cyclosorus but differs in this gradual tapering to very small pinnae on the lower blades. Locally common along streams at elevations of 100–2,200 m on all of the main islands except Ni‘ihau and Kaho‘olawe. Listed by Palmer (2003) as Amauropelta globulifera.

Woodsiaceae

(LADYFERN FAMILY)

Terrestrial ferns. Key feature is the presence of 2 ribbon-shaped vascular bundles seen in a cross section of the base of the stipe. Fronds are erect and 1–4 pinnate. Sori abaxial, round, “J” shaped, or linear in shape with kidney-shaped to linear indusia or occasionally without indusia. Virtually cosmopolitan in distribution. There are 15 genera and about 700 species, with 85% of these in the genera Athyrium and Diplazium. Sometimes the family is designated as the Athyriaceae and at

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other times merged into a broad concept of the Dryopteridaceae. Represented in the Hawaiian Islands by 4 indigenous genera. Athyrium is cultivated.

Athyrium—A cosmopolitan genus of ferns

that are terrestrial or growing on rock. The genus is found primarily in the north temperate regions but extends widely through the tropics as well. The fronds are usually large, 1–3 pinnate, with round or usually somewhat elongate sori covered by an indusium (rarely without). There are about 180 species. The name is from the Greek athyros, doorless, referring to the sporangia, which slowly push back the outer edge of the indusium. In the Hawaiian Islands represented by 1 endemic species.

Athyrium microphyllum (Sm.) Alston ‘ākōlea

Cystopteris—A cosmopolitan genus of 12

fern species distributed in both temperate and tropical habitats. The sori are usually rounded and are covered by an inflated, bladder-shaped indusium, which gives rise to the common name of bladder ferns. The name comes from the Greek kystos, bladder or sac, and pteris, fern. Represented in the Hawaiian Islands by 2 endemic species. Palmer (2003) has suggested that plants growing on wet ledges of the summit region of Mauna Kea and the subalpine zone of Haleakalā may be undescribed taxa. Cystopteris fragilis grows to high alpine elevations in North America. Cystopteris douglasii occurs on Maui and Hawai‘i, while Cystopteris sandwicensis occurs on Kaua‘i, O‘ahu, Lāna‘i, and Maui.

Deparia prolifera (Kaulf.) Hook. & Grev. ENDEMIC | SECURE

Cystopteris douglasii Hook ENDEMIC | RARE

ENDEMIC | SECURE

5.115 

5.114 

Cystopteris douglasii

Small to medium-sized terrestrial ferns. Blades ovate-lanceolate, obtuse tipped, 10–30 cm in length, pinnate-pinnatifid, tips of the pinnae obtuse. Sori position midway between margin and midrib. Rare on wet ledges in mesic forests and cave mouths at elevations of 1,500–3,000 m or more on East Maui and Hawai‘i. 5.113 

Athyrium microphyllum

Medium-sized terrestrial ferns. Rhizome forming a stout, erect caudex 15–30 cm in height. Fronds 25–90 cm in length, arrayed in rosettes at the tip of the caudex; blades dark green, 3 pinnate-pinnatifid to 4 pinnate and overall lacy in appearance; costae bearing short white translucent spines. Locally common in mesic to wet forests at 500–2,320 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe.

Deparia—A genus of about 50 small to medium-sized terrestrial ferns with a paleotropical distribution, with 1 species in North America. Blades pinnate to bipinnate. Sori generally linear along the veins and sometimes paired on both sides of the veins. The name comes from the Greek depas, which is a cup or a beaker, describing the shape of the indusium. In the Hawaiian Islands represented by 5 endemic species and 1 additional naturalized species.

Deparia prolifera

Medium-sized terrestrial ferns. Rhizomes trailing to erect. Fronds 35–120 cm in length; blades 1 pinnate-pinnatidfid to 1 pinnate-pinnatisect, linear triangular in shape, glabrous; only minimally narrowed at the base. Sori marginal and extending the margins of the pinnae. Nearly always with vegetative plantlets on the upper rachis. The base of the stipe generally without prominent scales. Locally common in wet forests at elevations of 215–1,280 m on all of the main islands except Ni‘ihau and Kaho‘olawe.

Diplazium—A genus of about 300–400 mostly terrestrial ferns found throughout tropical regions with a few extending into north temperate latitudes. The fronds are usually large, borne in clusters, and have simple to 3 pinnate-pinnatifid blades. The spores usually have prominent wingshaped folds. The name is from the Greek di, two, and plazion, oblong, referring to the double sori. In the Hawaiian Islands represented by 3 endemic and 1 naturalized species.

FERNS AND FERN ALLIES

Diplazium sandwichianum (C. Presl) Diels hō‘i‘o, pōhole ENDEMIC | SECURE

5.116 

Diplazium sandwichianum

Large terrestrial colony-forming ferns. Fronds up to 150 cm in length; stipes 45–90 cm long; blades 2 pinnate-pinnatisect to 3-pinnate-pinnatifid, deltate-ovate to ovate in shape, medium to light green, glabrous, chartaceous. Sori arranged in parallel rows along the midrib of the pinnae. Common and often forming a dominant ground cover in mesic to wet forest at 105–1,850 m elevation on all of the main islands except Ni‘ihau and Kaho‘olawe. Hawaiians still today gather and eat the raw fiddleheads and young fronds.

Vittariaceae—Recent molecular studies now place this family within the Pteridaceae.

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Colocasia esculenta

CHAPTER 6

Polynesian Introductions H

voyaging canoes appear to have also been naturally present in the Hawaiian Islands before human settlement. These are Pandanus tectorius (hala, screwpine) and Cordia subcordata (kou). The coastal tree Thespesia populnea (milo) may also fit this category, as it was an important tree to the Polynesians for its shade and for a variety of uses of its wood and fiber for rope. Hibiscus tiliaceum (hau) has also been questioned as to whether it was naturally present or first introduced by the Polynesians. The narrow ecological niche that this species occupies in the Hawaiian Islands and the complete lack of any pollen or seeds or any other subfossils in prehuman levels suggest it is in fact most likely a Polynesian introduction. The 25 plant species described here include 24 species shown quite conclusively by research to be Polynesian introductions, along with Schizostachyum glaucifolium (‘ohe, bamboo), which is a likely Polynesian introduction, though its status is still questionable. It should also come as no surprise that the Polynesians appear to have inadvertently introduced a number of weedy alien species to the Hawaiian Islands. These include Ludwigia octovalis (kāmole, primrose willow), Oxalis corniculata (‘ihi ‘ai, yellow wood sorrel), and Digitaria setigera (kūkaepua‘a, itchy crabgrass).

istorical evidence suggests that Polynesian ancestors moving eastward from Melanesia had colonized Fiji, Samoa, and Tonga by about 900 BC. For unknown reasons, Polynesian expansion from the homeland areas into Eastern Polynesia stalled for 500–1,000 years. New research has dramatically altered settlement dates, suggesting that Polynesians colonized the central Society Islands between AD 1025 and 1120 and dispersed to New Zealand, Hawai‘i, Rapa Nui, and other locations between AD 1190 and 1290. The date for the first arrival of Polynesians to the Hawaiian Islands has remained controversial, with suggestions for an early arrival from the Marquesas about AD 400–500 followed by a major colonization in the thirteenth century from Raiatea and Bora-Bora. The most recent analyses based on carbon dates have found no evidence of the earlier colonization and a more refined date of 1220–1261 for the first Polynesian arrival in the Hawaiian Islands. The exploration and settlement of eastern Polynesia involved the need for increasingly greater voyaging distances and thus more sturdy voyaging canoes built with double hulls. These large vessels had the advantage of sufficient space to hold multiple families, domestic goods, animals, and horticultural stock of useful plants necessary for long voyages of colonization. However, the size of these oceangoing canoes meant that they required superior sails for propulsion and more advanced experience with navigation. Although the early Polynesian voyagers were skilled navigators, most archeologists feel that the primary motivation for their exploratory voyages was to ease the demands of burgeoning populations. Polynesian mythology speaks of heroic discoverers of new lands for the benefit of those who voyaged on the expeditions rather than of explorers bent on conquest of new territories. Whatever the history of settlement, later Polynesian colonists introduced a line of high chiefs, the kapu system, the practice of human sacrifice, and the building of heiau. The ancient Polynesian settlers of the Hawaiian Islands brought many plant species with them on their voyaging canoes, and these plants were important for many uses in Native Hawaiian culture. In addition, they brought domestic pigs, chickens, and dogs on their voyaging canoes, as well as Polynesian rats. The precise number of plant species brought deliberately to the Hawaiian Islands by the early Polynesian settlers remains uncertain. The accepted number is 25–31 species, but there are conflicting opinions about several of these. At least 2 species that were almost certainly carried by the Polynesians on their

6.1  Ulu from Kanehunamoku, a block print by Dietrich Varez. Kanehunamoku is a mythical floating island. Two fishermen lost at sea came upon this land of abundance. Here, they discovered breadfruit (‘ulu) and brought it to Hawai‘i.

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Food Plants From early times, kalo was the primary food of the Hawaiian people, supplemented by other principal and traditional foods, most notably breadfruit (‘ulu), sweet potato (‘uala), yams (uhi, pi‘a), greens, ferns, fruit, fish, and seaweed (limu).

Araceae Colocasia esculenta kalo, taro

6.2 

Colocasia esculenta

Kalo is a large herb up to 1 m tall that is widely planted around the tropics of the world today and is thought to be native to India. It displays large heart- or arrowshaped peltate leaves that are green above with dark purple to yellow venation beneath and arise from underground corms. Kalo continues to be cultivated in the Hawaiian Islands and persists outside of cultivation on all of the main islands except Kaho‘olawe. Kalo was the chief source of starch for the Hawaiians, who mashed the baked or steamed corms to make poi, which was believed to have the greatest life force of all foods. This gray, pasty gruel remains widely available today in restaurants and markets. The leaves, petioles, and spadices subtending the inflorescence of taro plants were cooked as greens. The cooked corm could also be sliced and eaten or grated and mixed with sweeteners and coconut milk and baked into a pudding called kūlolo. Cooking was necessary to break down the calcium oxalate crystals in the tissues. Kalo was also employed as a medicine to heal sores and reduce the inflammation of insect bites, as bait for a fish called ‘ōpelu, as an adhesive to glue kapa pieces together, and in

religious rituals. The Native Hawaiians grew about 300 varieties of taro, with individual cultivars adapted to specific ecological conditions related to soil type, microclimate, and rainfall.

Alocasia macrorrhizos ‘ape, elephant ears ‘Ape, or elephant ears as it is sometimes called, is a large herb with broad shinygreen leaves up to 1 m or more in length and 60 cm in width. The thick trunk can reach 1.5–2.5 m tall and 8–10 cm in diameter. These plants are related to and resemble the kalo plant but can be readily distinguished by their upright leaves, compared to kalo leaves that point downward. As with kalo, the underground stem and corm are edible after lengthy cooking to break down the toxicity of calcium oxalate crystals, which can injure internal human tissues. ‘Ape was not a favored food of the Polynesians and was eaten only in times of famine when other food was scarce. An old Polynesian saying states, “The eater of ‘ape will have an itchy mouth.” The leaves of ‘ape, together with ti leaves, were used to wrap a fevered person for comfort because the bitter sap was thought to frighten away bad spirits. ‘Ape is commonly used today as a landscaping plant for its tropical appearance and ability to provide garden shade.

Convolvulaceae Ipomoea batatas ‘uala, sweet potato

6.3 

Ipomoea batatas

The sweet potato originated in the neotropics and was first domesticated there by Native Americans. Interestingly, several varieties of sweet potato were widely cultivated in the Hawaiian Islands at the time of Captain Cook’s arrival in 1778, and its widespread cultivation in the Pacific almost certainly predates the voyages of Columbus as well. Molecular data

confirm the close relationship of domestic sweet potato in the Andes with the Hawaiian cultivars. This distribution and other new findings are increasingly indicating that there was early Polynesian contact with the west coast of the Americas. Sweet potatoes are herbaceous vines with erect or trailing stems, often rooting at the nodes, which arise from underground tuberous roots. A tuberous root is a modified lateral root, enlarged to function as a storage organ. In contrast, a stem tuber such as a potato is a fleshy underground storage organ derived from stem tissue. The leaves are variable, 5–10 cm in length, and the flowers are purple to lavender or white in some varieties. The sweet potato is a member of the Convolvulaceae, or morning glory family, and is unrelated to yams, which are members of the Dioscoreaceae, a Monocot family. Sweet potato was a staple in the diet over broad areas of Polynesia. The Native Hawaiians are reported to have grown about 230 cultivars of the sweet potato, which varied in the growth conditions, sweetness, and starchiness. About 24 of these remain in wide cultivation today. To maximize the nutritional value, sweet potatoes were commonly baked in their scrubbed skins and then mashed to make sweet potato poi. They were also used as fish bait and to fatten pigs. In addition to the fleshy roots, the stem tips and young leaves were eaten. Mashed and fermented root tubers were used in the making of a beer. Special varieties of sweet potato had important medicinal properties. These were used as a tonic during pregnancy and to induce lactation, to cure asthma, as a laxative, and as a gargle for a sore throat. Raw sweet potato mixed with ti stem was used when necessary to induce vomiting. Kamapua‘a is said to be the god of the sweet potato. This god has a piglike snout, making it possible for him to excavate the large root tubers.

Ipomoea cairica koali‘ai, morning glory The origin of this cultivated morning glory is uncertain, as it is widely distributed around the world today in tropical and warm temperate latitudes. It is thought to have been cultivated by the early Polynesians for centuries. It is not clear whether this species represents a Polynesian introduction or alternatively

P O LY N E S I A N I N T R O D U C T I O N S

was present in the Hawaiian Islands by natural colonization. These two possibilities may both be true. Today it is sometimes cultivated as a garden plant, but it is known more as an aggressive invasive species. The growth rate of its twining herbaceous stems is extremely rapid and has given rise to the alternate name, “mile-a-minute vine.” The ovate to orbicular leaves are palmately divided into 5–7 lobes. Flowers may be bluish, purple, or white with a purple center. The Hawaiians used its tuberous roots as famine food.

as a food except in times of famine. The pi‘a yam differs from the uhi yam in having round, wingless stems rather than square vine stems with wings. The palmate leaves are divided into 3–5 lobes instead of being ovate as in uhi.

Moraceae Artocarpus altilis ‘ulu, breadfruit

Dioscoreaceae Dioscorea alata uhi, yam Uhi is a trailing herbaceous vine growing up from large starchy tubers and one of 3 yam species thought to have been brought to the Hawaiian Islands by the early Polynesians. The edible tubers of uhi were cooked in an underground oven and then eaten warm, as with other edible yams. In cultivation these were grown over frames or supports, but they trail over shrubs and trees when naturalized. Although uhi was a staple food in other parts of Polynesia, the Native Hawaiians considered it inferior to other starchy tubers for making poi. Yams should not be confused with ‘uala, sweet potatoes.

Dioscorea bulbifera hoi, bitter yam Bitter yam is a trailing herbaceous vine with long twining stems that can extend for tens of meters across the ground surface. This growth habitat allows plants to become aggressive invaders in favorable habitats. It produces fleshy root tubers from the axils of its large heart-shaped leaves, a condition giving rise to the name “air potato.” These bitter yams are poisonous unless rinsed in running water for several hours prior to cooking and thus were used primarily in times of famine.

Dioscorea pentaphylla pi‘a, wild yam This wild yam is another herbaceous vine with large fleshy root tubers and is a close relative of uhi. It was grown in wet valleys on the windward slopes of the major islands, but it likely had only limited use

to termites and shipworms, making it a valuable timber for surfboards, structures, and outrigger canoes. The milky sap was useful as glue and caulking material in canoe building. Another use of the sap was to make a birdlime used to catch nectar-feeding birds, which were highly valued for their brightly colored feathers. The wood pulp of breadfruit can be used to make a paper called breadfruit tapa. It was also used in traditional medicine to treat a variety of illnesses ranging from sore eyes to sciatica. Today, breadfruit is widely planted as a shade tree in home gardens and as a source of its fruit.

Musaceae Musa x paradisiaca mai‘a, banana

6.4 

Artocarpus altilis

Breadfruit is a large tree, 12–20 m or more tall, with a broad crown. The existing genetic forms have an ancient origin, with hundreds of known cultivars, but all are likely native to Melanesia. The large entire to deeply lobed leaves, up to 1 m in length, are iconic silhouettes in many Hawaiian quilts. The large globular breadfruit, up to 20 cm or more in diameter and weighing up to 10 kilograms (kg), are composites of thousands of small fleshy fruits growing together. The flesh is a mealy sweet pulp, similar in many ways to potato. The fruit is usually picked while still firm and softens in a few days. The starchy composite fruits are roasted, baked, fried, or boiled. When cooked, the taste is potato-like or reminiscent of fresh baked bread, hence the name. Breadfruit was an important staple across many parts of the Pacific Islands because of the large yield of individual fruits, up to 100– 200 from a single tree. However, breadfruit was much less a staple of diet in the Hawaiian Islands compared to other Polynesian islands. Beyond its importance as a food source, breadfruit had many other uses. Its lightweight wood is resistant

Bananas are tall herbs up to 10 m tall, with large leaves spirally arranged toward the ends of what is called a pseudostem. The inflorescences are large, divergent to pendent, bearing many staminate and fewer pistillate flowers that produce a jellylike nectar. In the Hawaiian Islands, bananas are sparingly surviving from early plantings in mesic to wet forest on all of the main islands except Ni‘ihau and Kaho‘olawe. There are hundreds of cultivars of the banana, and about 70 of them were extensively cultivated by the Hawaiians. The plant had uses beyond that as a food source. The stalk, leaves, and fruit were also used in religious ceremonies. The stalks were employed to line underground ovens, as canoe rollers, and mashed to make a poultice for sprains and broken bones. Parts of the leaves were used in lei making, and the sap had value as a dye. Nectar from the flowers was used as a baby food.

Musa troglodytarum mai‘a he‘i, mai‘a polapola, banana This banana species is thought to be a relatively recent Polynesian introduction from Tahiti. It is also a tall herbaceous perennial reaching up to 12 m tall. It served as an important carbohydrate crop. The bananas were cooked when mature to produce a flavor similar to breadfruit. The black stems were used in weaving with Pandanus. It was once widely cultivated and persists sparingly in low-elevation mesic to wet valleys on most of the main islands.

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Myrtaceae

Taccaceae

Syzygium malaccense ‘o‘hia ‘ai, mountain apple

Tacca leontopetaloides pia, Polynesian arrowroot

Mountain apples are small trees, up to about 15 m tall, that can be found widely planted in wet forest areas at lower elevations on all of the major islands. They are members of the Myrtaceae or eucalyptus family, with an origin thought to be in Southeast Asia. The red, pink, or white oval fruit, 5–8 cm in diameter, is highly prized for its sweet and juicy flesh. The delicate taste is not very distinctive but somewhat like a pear. The trunks of these trees were used as house beams. Extracts from the bark were used to cleanse minor skin wounds.

Pia is a perennial herb arising from fleshy and starchy tubers that are replaced each year by new ones that develop from a thick, downward-growing rhizome. It is a member of the Taccaceae, a small family related to the Dioscoreaceae. Because of its extremely widespread planting in ancient times, the natural range of Polynesian arrowroot is not entirely clear. Wild-growing plants can be found from Africa and Madagascar across India to Southeast Asia, Malesia, and northern Australia on into the Pacific, where it was carried by the Polynesian voyagers. It is cultivated today in portions of Polynesia and Southeast Asia, but cultivation in the Hawaiian Islands largely ceased when Euro-­ American colonists brought alternate carbohydrate crops. Polynesian arrowroot can be occasionally found today growing in former Polynesian living sites on Kaua‘i, O‘ahu, Moloka‘i, Maui, and Hawai‘i. In cultivation, tubers were planted in small holes in the ground at the end of the rainy season and required little care if adequate water and fertile soil were provided. The above-ground tissues of the plants die back at the end of summer, and the tubers are harvested in winter when the plants are dormant. The starch obtained by cooking the tubers was used as a thickening agent, much as cornstarch, and often mixed with coconut milk. Because of bitterness caused by the presence of a chemical called taccalin, Polynesian arrowroot was much more a famine food than a major part of regular Hawaiian diets. The plant also had medicinal value in treating diarrhea and dysentery.

Poaceae Saccharum officinarum ‘kō, sugarcane

6.5 

Saccharum officinarum

Sugarcane is a perennial grass widely cultivated at low to moderate elevations. Its probable origin is in Southeast Asia. The unbranched stems or stalks reach heights of 2–6 m and are 2–5 cm in diameter, with clearly marked internodes much as in bamboo. In addition to their use as a sweetener, tall hedges of sugarcane provide an effective windbreak, and the foliage can be used as thatch. There is some controversy over whether or not sugarcane was brought by the early Polynesian voyages or was instead a later naturalization. In earlier times, the people of Hawai‘i grew as many as 40 different varieties of sugarcane.

Kava grows as a shrub up to 3 m tall, with jointed stems that are swollen at the nodes and heart-shaped leaves. Plants dioecious. Kava, or ‘awa as it is called in Hawaiian, is a member of the Piperaceae, or pepper family. Its origin is uncertain, but it was widely spread by early human migrations throughout the Pacific Islands, and multiple cultivars exist. In the Hawaiian Islands, kava is cultivated and persists in shaded mesic valleys or windward mesic sites on all of the main islands. The name applies to both the plant and a drink made from the plant. Kava has a sedating effect and is primarily consumed to relax without disrupting mental clarity. Its active ingredients are called kavalactones. Today, kava extract is marketed as herbal medicine to treat stress, insomnia, and anxiety. The fresh or dried root and lower stem parts were used in making a special drink for the Hawaiian aristocracy (ali‘i). This strongly bitter drink was important in religious and social ceremonies because of its narcotic properties. With increasing popularity throughout the Western world (kava bars) as well as in Polynesia, concern has been raised about potential effects of kava in causing liver damage.

Utilitarian Plants Euphorbiaceae Aleurites moluccana kukui, candlenut

Aleurites moluccana

Piperaceae

6.7 

Piper methysticum ‘awa, kava

Kukui are distinctive, large, spreading trees 10–20 m tall and up to 90 cm in diameter, with large 3–5-lobed leaves having a distinctive color. These trees, native to Malesia, form a conspicuous element of mesic valleys and ravines and are easily recognized by their pale foliage, which stands out next to the darker color of other trees and shrubs. The plant had many uses for the Hawaiians. The round, greenish nuts contain 1–2 seeds with a very hard seed coat and high oil content. The nuts were

6.6 

Piper methysticum

P O LY N E S I A N I N T R O D U C T I O N S

strung on palm-leaf midribs and used for torches. The oil from the nuts was used in lamps, and the juice of the husk produced a black dye. The nuts were also used as a laxative. The tree trunks were employed in canoe construction, while gums and resins had medicinal properties. During the nineteenth century, kukui oil was exported to Asia, where it was used as a drying oil in paints and varnishes. A fungus was grown on the decaying wood of the trees and exported to China for local consumption. Today the nuts are commonly polished and strung into lei for tourists. The kukui is the official state tree of the Hawaiian Islands.

paper mulberry has become an aggressive invasive species in many areas where it has been introduced, particularly in Central America, South Asia, and portions of the southeastern United States. The trees rapidly reproduce vegetatively from root suckers, creating thickets of scrubby stems.

Clusiaceae

immediate coast are famous for virtually horizontal trunks that extend out over the water.

Arecaceae Cocos nucifera niu, coconut palm

Calophyllum inophyllum kamani

Moraceae Broussonetia papyrifera po‘a‘aha, wauke, paper mulberry

6.8 

Broussonetia papyrifera

Paper mulberry is a small semideciduous tree reaching up to 15 m tall, with broad leaves of variable size and shape. The leaves can be ovoid to deeply lobed and 7–20 cm in length, with a rough surface above and fuzzy hairs below. It is native to eastern Asia, with a range from India through China to Japan and south to Indonesia. There are separate male and female flowers, and the female flowers produce orange to red fleshy fruit 3–4 cm in diameter. The fruit is highly edible and an important food for wildlife, but it is too fragile to be commercialized for human use. The significance of paper mulberry to the Polynesians came from its use as the preferred plant for making soft and durable tapa or kapa cloth. The inner bark contains strong fibers that can be washed and pounded to produce a high-quality cloth that was much prized. The traditional art of making kapa cloth declined precipitously with the introduction of imported fabrics, but it has made a resurgence in recent years with increased interest in traditional Hawaiian crafts. Although it has been less of an issue in the Hawaiian Islands, fast-growing

6.10  6.9 

Calophyllum inophyllum

Kamani is a large tree reaching 20 m tall, with a rounded canopy of dense foliage and a trunk that typically leans away from an upright position. The primary Polynesian use of kamani was its reddish wood, which was carved into canoes, used as timber for buildings, and in particular for making wooden containers and bowls. Along with wood from Thespesia and Cordia, kamani was particularly prized for food dishes and food storage vessels, as the wood imparted no unpleasant odor or taste to the food. It was widely cultivated in the area around houses in Hawaiian villages, as well as in groves near temples. All parts of the kamani were used by the Polynesians. Flowers were used in lei making and placed in kapa storage bags to give them a sweet fragrance. The mature globular fruit is about 5 cm in diameter, with a leathery skin covering a corky mass of storage tissue and an oil-rich seed. Although the seeds are poisonous, a thick oil was extracted from them and sometimes used in place of kukui oil for lamps. The oil, enhanced with coconut oil and flower fragrances, is used for traditional lomi lomi massage. Kamani seeds were also polished to give them a look similar to kukui and used in lei, earrings, and other jewelry. These slow-growing trees are well adapted for sandy soils and coastal sites exposed to aerosol salts. Today kamani is grown in coastal and other open sunny areas and is used as a shade and street tree. Naturalized trees growing along the

Cocos nucifera

Coconut palms are trees with solitary trunks up to 30 m tall and pinnately compound fronds up to 6 m long. The green or yellowish fruit houses a large hollow seed that contains a large quantity of liquid endosperm or coconut water. Coconut palms are widely cultivated in the Hawaiian Islands but sparingly naturalized and were usually found in areas of Polynesian habitation. Although brought by the Polynesians, coconut palms were never used as extensively in the Hawaiian Islands as in other parts of Polynesia. Coconut palms are present on all of the main islands and Laysan in the Northwestern Islands. Two forms of this plant were introduced by the Polynesians. One form, niu hiwa, has dark green fruit and a black shell when mature. It was used in cooking, medicine, and in religious ceremonies. Another form, niu lela, has a reddish yellow exterior and yellowish shell when mature. It was used for more secular purposes, with the trunks used as house posts, for canoes, and as food containers and drums. The palm fronds were employed in basket weaving and for thatch. The leaflets were made into fans, while the midribs were used as brooms, to string kukui nuts, and for other uses. The husks of the fruit were burned as fuel and the separated fibers were made into strainers for kava and other liquids and to make sennit, a braided cordage used in the construction of boats, houses, and other structures. The shell of the fruit was used for scooping, eating, and drinking. The water from the fruit was a favored

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beverage, and the nuts were a water source for long journeys. The flesh of the fruit was an important food source in all stages of maturity. Oil from the mature flesh was rubbed on the body and hair. Today, coconut palms are extremely common along Hawaiian beaches and streets and are widely used in landscaping. Despite the nostalgia for coconut palms as emblematic of tropical coasts, the trees often present a liability because of problems and dangers from falling fronds and coconuts.

Boraginaceae Cordia subcordata kou Kou are small trees, sometimes forming thickets, with thin, leathery leaves and wrinkled orange flowers. Kou is a member of the Boraginaceae, or borage family. It is thought to be native to the Malesian region but now occurs throughout the Pacific due to early human movements. Although almost certainly brought to the Hawaiian Islands by the Polynesians on their voyaging canoes, fossil evidence has now established that kou had already colonized the Hawaiian Islands naturally before their arrival, and it is now considered to be an indigenous species. In the Hawaiian Islands, kou is found today in low-elevation, dry coastal areas on all of the islands except Moloka‘i and Kaho‘olawe. The wood was used to make bowls and utensils. The flowers were made into lei and the white seeds consumed. Leaves were also used to color fish lines. Along with kamani, kou was planted near temples.

Asparagaceae Cordyline fruticosa ti, kī

6.11 

Cordyline fruticosa

Ti plants are shrubs 3–5 m tall with unbranched or few-branched stems. The thin leaves are spirally arranged toward the ends of the branches. This species is now usually placed in the Asparagaceae, but older books identify it as Cordyline terminalis and place it in the Agavaceae. The origins of ti are obscure because it has been cultivated for thousands of years, but the likely guess is that it originated in Southeast Asia and Malesia. From there it was widely spread across the Pacific by early Polynesian migrations. The ti plant was considered sacred to the Hawaiian god Lono and to the goddess of the hula, Laka. It was also an emblem of high rank and divine power. The Hawai‘i feather kāhili is thought to have been inspired by this plant. The leaves were also worn or carried as protection to ward off so-called evil spirits and to call in good spirits. The kahuna (priests) conducting ceremonial rituals wore ti leaves either as a lei or as garments. The leaves had a broad variety of everyday uses, including wrappings for food to be cooked or preserved, eating plates, thatching for roofs, insect fans, and wearing apparel such as rain capes, hula skirts, and sandals. The roots were baked and either eaten or used to make an alcoholic beverage (‘ōkolehao). In the Hawaiian Islands today, ti plants are widely present in mesic valleys and forest areas, as well as extensively cultivated as hedges, windbreaks, and garden plants on all of the main islands except Kaho‘olawe. New cultivars have produced foliage in a variety of colors.

Zingiber zerumbet ‘awapuhi, wild ginger Wild ginger is a tall herb growing from creeping aromatic rhizomes. The numerous distichous leaves have swollen petioles. The flowers are borne in terminal, ovoid, globose, or cylindrical spikes that arise on leafless shoots. Wild ginger is thought to be native to India but has been widely carried by humans throughout the Pacific Islands. In the Hawaiian Islands, wild ginger is naturalized and common in disturbed, mesic shaded places on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Maui. The fluid from the inflorescence was used as a shampoo, the powdered rhizome for scenting kapa, and the leaves as a flavor enhancer for meat. Ash from burned leaves was used medicinally to reduce inflammation and related pain and to treat worm infestation and diarrhea.

Malvaceae Hibiscus tiliaceus hau

Zingiberaceae Curcuma longa ‘ōlena, turmeric ‘Ōlena is a herbaceous perennial plant in the Zingiberaceae, or ginger family, with a short, upright leafy stem arising from a thick underground rhizome below. The rhizome is powdered to produce turmeric, widely adopted in many cuisines. A major use to the early Polynesians, however, was the grinding of the rhizome to yield a yellow to orange dye used in kapa cloth. Another traditional use for ‘ōlena was as a spiritual purifier with great power. Pieces of the powdered rhizome were mixed with seawater and sprinkled onto an ill person or newly built structure, along with appropriate prayers, as a blessing to remove negative forces.

6.12 

Hibiscus tiliaceus

Hau is a pantropical polymorphic species sometimes placed in a segregate genus as Talipariti tiliaceum. This arboreal form of hibiscus is widely distributed in tropical and subtropical regions of the world, and its long history of human use and widespread distribution of the genus Hibiscus makes it very difficult to determine its native range. It is abundant in the Hawaiian Islands today along freshwater streambeds, brackish coastal marshes, and moist gullies where it grows in dense thickets on all of the main islands except Ni‘ihau and Kaho‘olawe, as well as on Midway Atoll and French Frigate Shoals.

P O LY N E S I A N I N T R O D U C T I O N S

As mentioned above, the early Polynesians were certainly aware of hau and likely brought it with them to the Hawaiian Islands because of its many uses. Generally hau grows as a shrub or small tree from 2 to 10 m or more tall, with many sprawling branches that form an impenetrable tangle. The heart-shaped leaves are smooth above and velvety beneath. The flowers open yellow, fading to orange and finally dark red by the end of the day as the pigments oxidize. In the Hawaiian Islands, hau is common along streams, river mouths, and coastal areas at elevations from sea level to 300 m or more. The Native Hawaiians had many uses for hau. The stem fibers were used for cordage and the light wood for the spars of outrigger canoes, for floats for fishnets, and for tinder to start fires. The sap had medicinal value to relieve chest congestion, and the flowers and foliage were used in lei making as part of traditional hula ceremonies.

Cucurbitaceae Lagenaria siceraria ipu, gourd

Cucurbitaceae, or squash family. Gourds for household use were prepared from the mature fruit by first cutting off the top and filling the gourd with seawater, which was changed every ten days to soften the gourd’s hard flesh. At this point the flesh was then removed along with the seeds. Coral fragments were used to smooth the roughness of the inside of the dried gourd rind. A shell or carved wood was then fashioned to create a stopper that fit into the gourd’s opening. Depending on their quality and shape, gourds could be used for carrying food or water. Ipu were also important for the construction of rattles and drums for hula and chanting, and they are still considered essential today for those purposes. Ancient Hawaiians grew gourds primarily on the southern and leeward coasts of the Islands, as the plants favor hot and sunny habitats with plenty of moisture. Generally, the gourds were planted at the start of the rainy season and matured over the summer to be ready for a fall harvest in about six months.

Rubiaceae Morinda citrifolia noni

paired, except where forming fruit. Thick and oval in shape, these leaves are deep veined, short stemmed, and 20 cm or more in length. The globose heads of small white flowers mature to form a multifaceted fruit 7–10 cm in diameter. The fruit resembles small versions of breadfruit and is formed throughout the year. Although the fruit has a strong unpleasant odor and taste, it is still widely used medicinally. The fruit and its juices have been used in the treatment of diabetes, heart troubles, high blood pressure, urinary disorders, skin rashes, and joint pain. Dyes for kapa cloth were extracted from the bark and roots.

Poaceae Schizostachyum glaucifolium ‘ohe, Hawaiian bamboo ‘Ohe is a clumping bamboo that when mature can reach up to 12 m or more in height, with stems up to 8 cm in diameter. As with all bamboos, it is a member of the Poaceae or grass family. This species is thought to have its origin in India or perhaps Indonesia and was brought very early to the Hawaiian Islands. Lengths of hollow stems and closed ends were used as water containers, along with gourds. Bamboo stems had many uses: building posts, bridges, irrigation channels, floats, masts, furniture, utensils, agricultural tools, ladders, ornaments, toys, musical instruments, and fishing poles. Split bamboo was woven into wall and sleeping mats, screens, baskets, fans, and roof thatch.

Fabaceae Tephrosia purpurea ‘auhuhu, fish poison plant

6.13 

Lagenaria siceraria

Ipu is a spreading vine with branched tendrils and broad, heart-shaped leaves 10–15 cm in diameter. The fruit of the ipu, which varies greatly in size and shape, becomes tan or beige in color with maturity. Gourds are members of the

6.14 

Morinda citrifolia

Noni is an evergreen shrub or small tree, usually less than 3 m high but occasionally as tall as 6 m, in the Rubiaceae or coffee family. The conspicuous, large, dark green leaves are shiny and generally

‘Auhuhu is a small shrubby legume that is widespread in distribution today across the paleotropics. It reaches heights up to 1.5 m, with compound leaves of 7–21 leaflets and large pink to white-purplish flowers. The elongate seed pods were pounded by the Polynesians to extract a chemical compound called tephrosin. Placed in tidal pools, tephrosin stuns fish and causes them to float to the surface where they can be easily caught. Tephrosin was also used as an external ointment to treat skin diseases.

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Spathodea campanulata

CHAPTER 7

Alien Invaders

I

today. In many cases these naturalized plants are restricted to disturbed areas or roadsides and are not able to invade native plant communities. Unfortunately, more than 100 species within this group have become aggressive invaders, readily able to become established in native habitats, where they have the potential to cause serious environmental impacts. These species can fundamentally change ecosystem structures and processes by crowding out or shading native species, enriching soil fertility and thus facilitating alien grass invasion, and altering natural fire cycles.

n a sense, all plants in the Hawaiian Islands are the product of alien invaders, plants whose ancestors were able to disperse over thousands of kilometers of ocean to reach the Islands. This process of natural dispersal and successful establishment occurred over millions of years without human assistance. Over the time scale of centuries, however, mankind has added greatly to the number of arrivals. One group of such nonnative or alien species arrived with the Polynesians, as described in chapter 6, who brought a variety of cultivated plants with them. Only a few of these have escaped from cultivation and become a part of natural plant communities today. Nevertheless, the Native Hawaiians did have a major impact on the lowlands of the Islands that greatly facilitated later invasions. This impact came from widespread clearing and burning of the native vegetation. Other species widespread today appear to have come inadvertently with the Polynesians as weedy invaders. Beginning with the arrival of Captain Cook in 1778, the rate of plant introductions increased and has continued to increase almost exponentially with a constant influx of humans and cargo from all over the world. Each new cultural group arrived with their favorite decorative garden species, medicinal plants, and plants special for their cuisine. Agricultural interests accelerated the process more by bringing in pasture grasses and herb seeds to support grazing animals. Often, their sources of seed were contaminated with weeds from other sources. Finally, foresters unappreciative of native tree species introduced a range of alien trees to reforest watershed areas decimated by cattle grazing, deforested, or opened up after ‘ōhi‘a dieback events. Introductions of tropical plant species of horticultural interest continues today. The recently published Tropical Garden Flora by George Staples and Derral Herbst (Bishop Museum Press, 2005) describes 8,134 introduced species, and it is likely no exaggeration to say that more than 10,000 species of plants have been introduced to the Hawaiian Islands for garden cultivation or other uses. Many of these alien species have very specific habitat requirements and remain in the gardens, parks, and rangelands where they were planted. However, a subset of this large pool of introduced species has been able to disperse their propagules and become established away from cultivation, becoming what we term naturalized species. Wagner et al. (1990) in their Manual of the Flowering Plants of Hawai‘i listed 861 species as naturalized, and a 1999 update to the flora raised this number to 1,044. Adding in the alien ferns that were not included and newly naturalized species, this number is likely more than 1,100 naturalized species

Lowland Dry Forests and Shrublands The driest areas of lowland in the Hawaiian Islands, with elevations below 300 m and less than 500 mm of rain each year, are now almost totally dominated by alien plant species. These areas that once supported arid grasslands and shrublands were the principal habitation of the aboriginal Hawaiians and were greatly disturbed by clearance, fire, and erosion. Native species today are common in only a few of the driest coastal areas of the Islands, as, for example, on Diamond Head and Ka‘ena Point on O‘ahu. The introduction of Leucaena leucocephala (koa haole; Fabaceae) by European colonists and its subsequent spread resulted in its rapid colonization and ultimate domination of these areas. It was one of the earliest alien plants, likely introduced by European colonists to the Hawaiian Islands in the early nineteenth century. This spineless legume shrub was widely planted to provide a fast-growing source of firewood, charcoal, and foliage for animal forage. The species was considered a potential “miracle tree” for planting in the dry tropics of Asia because of its drought tolerance and nitrogen-fixing root nodules that allow it to grow rapidly in arid, poor soils unsuitable for crop plants. However, in the Hawaiian Islands it has crowded out the natural plant communities at low elevations and permanently altered soil conditions, making it difficult for native species to become established. In its early introduction, grazing pressure kept Leucaena from becoming a dominant landscape species until cattle numbers were sharply reduced about 1920. Without this controlling element, Leucaena spread rapidly, and its coverage expanded before World War II with aerial seeding over broad areas to control erosion. Today, Leucaena scrub blankets the lowlands of O‘ahu and Kaua‘i and covers broad lowland areas on all of the major islands.

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7.1 

Leucaena leucocephala

Prosopis pallida (kiawe; Fabaceae), a native of the neotropics, was introduced to the Hawaiian Islands in 1828 by a French priest who received seed from the Royal Botanic Garden in Paris. The pods of this species are a highly desirable food for cattle because of their high protein content. Although deliberate planting of this tree was quickly accomplished, its distribution expanded dramatically from the germination of seeds passing through the digestive system of cattle. In this way, kiawe became widely established everywhere in coastal flats and lowlands where cattle walked. Its deep roots readily tap into lens of freshwater along the coast, producing open stands and savannas of kiawe, with individual trees reaching as tall as 10 m. Without control by cattle grazing, Prosopis has expanded greatly to form impenetrable spiny thickets in coastal dryland areas. Much of the area dominated today by kiawe was likely once native dry forest.

known as Acacia farnesiana; Fabaceae) and Opuntia ficus-indica (pānini; Cactaceae) were widely naturalized in dry lowlands before koa haole and kiawe but have been replaced today over much of their range by the more aggressive koa haole. Vachellia farnesiana is a thorny, deciduous shrub that grows to 4 m tall, sometimes forming impenetrable thickets, although in most areas it forms a more open cover on rocky slopes. The seeds of klu are dispersed by grazing animals, which eat the pods. Although the aerial portions may be killed by fire, it soon regenerates from basal shoots. Vachellia farnesiana grows in dry habitats between sea level and 400 m elevation on all islands. There are dense infestations at Lualualei on O‘ahu and Līhau on Maui, as well as in some overgrazed areas on Lāna‘i and the Hā‘upu area of Kaua‘i. Opuntia ficus-indica was once an aggressive invader of pastures and dry rocky slopes, but its abundance has been greatly reduced by the introduction of biocontrol by a moth species, Cactoblasus cactorum.

7.3 

7.2 

Prosopis pallida

Two other early introductions to the Hawaiian Islands from Central America, Vachellia farnesiana (klu, formerly

Vachellia farnesiana

Lantana camara (lantana; Verbenaceae) was first introduced to the Hawaiian Islands as an ornamental in 1858 from the West Indies but quickly escaped to become naturalized throughout the Islands. This prickly shrub with strong-scented leaves rapidly became an aggressive invader in dry forest areas, as it has done in the dry tropics around the world. The small fleshy fruits are dispersed by alien fruit-eating birds and invade open pastures and dry forest habitats. Once established, Lantana forms dense shrubby thickets that choke out most native species. Unlike many native shrubs, Lantana

is capable of surviving all but the hottest fires, regenerating from basal shoots. Although not eliminating its presence, several biocontrol insects have been effective in reducing the impact of Lantana on native plant communities.

7.4 

Lantana camara

Casuarina equisetifolia (ironwood, beach she-oak; Casuarinaceae) is native from Australia eastward into Melanesia and north to Southeast Asia, but it was also an early human introduction over a much broader area. It was brought to the Hawaiian Islands in 1882 with plantings initially on Kaua‘i. Casuarina equisetifolia has long been considered a useful tree for windbreaks, erosion control, and firewood production, and it is thus a characteristic feature of coastal landscapes of tropical and subtropical countries around the world. This fast-growing tree with foliage resembling pine needles can reach a height of 20 m in as little as 10–12 years and an eventual height of 40 m or more. Its tolerance of moderately saline soils and the presence of nitrogen-fixing root nodules allow it to colonize poor soils. However, Casuarina has a strong tendency to become naturalized, as it produces thousands of wind-borne seeds and reproduces vegetatively through coppicing to produce dense stands that eliminate almost all other species. Once established, it radically alters the light, temperature, and soil chemistry regimes of coastal habitats, as it outcompetes and displaces native plant species. The ground below stands of these trees typically becomes ecologically barren. Unlike native coastal trees and shrubs, shallow roots make it susceptible to toppling during high winds, leading to increased beach and dune erosion. Casuarina is common today on all but the driest and wettest coastal areas of the islands up to 500 m elevation.

A L I E N I N VA D E R S

7.5 

Casuarina equisetifolia

Grevillea robusta (silk oak; Proteaceae), a large evergreen tree native to eastern Australia, was introduced into the Hawaiian Islands about 1880 as an ornamental and for its potential timber value. Its fernlike foliage and distinctive yellow-orange flowers have made it a popular street and park tree. It was also planted extensively in forestry reserves and is now naturalized widely on all of the major islands. Grevillea robusta is an aggressive invader of dry grassland areas and open lava fields up to 1,500 m elevation on all major islands, where it tolerates dry substrates with little soil development. It also grows well, however, under more mesic conditions. The winged seeds disperse broadly, often producing single-species stands. Because of its prolific reproduction, it has been classed a noxious weed on ranchlands on Hawai‘i. It is also a tremendous invader on Kaua‘i, visible just about everywhere on the road to Kōke‘e, as well as in the coffee-growing area above Kona, Hawai‘i. On Kaua‘i its spread was enhanced by damage from Hurricane ‘Iniki.

planted beginning about 1930 on O‘ahu, Maui, and Hawai‘i. Unlike most trees in the Hawaiian Islands, Citharexylum is deciduous during the dry season, and it can be readily recognized from a considerable distance when the leaves turn orange prior to their fall. Once thought to not readily naturalize, Citharexylum has escaped into low-elevation dry forests on O‘ahu and Maui. Alien fruit-eating birds are the dispersal agent for the fruit. A related species, C. caudatum, is naturalized on O‘ahu.

7.7 

Citharexylum spinosum

Coccinea grandis (ivy scarlet gourd; Cucurbitaceae) is a fast-growing vine first introduced to the Hawaiian Islands about 1969 as an ornamental and cultivated for its edible young shoots and fruit. The seeds are presumably spread by birds and perhaps rats and other small mammals, and Coccinea is now a widespread urban weed in the Hawaiian Islands. It is an aggressive vine that quickly covers nearby plants or structures such as fences and power lines. Because of its tolerance of dry, hot environments and its smothering habit, Coccinia has the potential to invade dry forest areas on all of the major islands.

Alien Grasses and Altered Fire Regimes

7.6 

Grevillea robusta

Although not yet a major problem, concern has been raised about Citharexylum spinosum (fiddlewood; Verbenaceae), a medium-sized tree that was widely

The introduction of alien grasses has changed much of the natural fire cycle in low-elevation dry forests and middle-­ elevation scrub habitats, as, for example, in Hawai‘i Volcanoes National Park. Much of the area in the park originally supported an open scrubby forest, separated by patches of sparsely vegetated lava fields. Fires, ignited by rare lightning or infrequent lava flows, were uncommon, but those that did occur remained small because of the natural vegetation mosaic. In the early 1970s, this situation changed

as the National Park Service began a management program to eliminate feral goats from the park. Two invasive alien grasses, Schizachyrium condensatum (beard grass) and Andropogon virginicus (broomsedge), had been held in check by the goats. With the removal of the goats, these two invaders quickly expanded, creating a continuous ground cover between islands of native scrub forest. The persistent flowering stalks of these species provide a flash fuel that readily carries fire across the landscape. These grasses are fire stimulated, rapidly resprouting from basal sprouts after burning. The total area burned by fires in lowland ecosystems of Hawai‘i Volcanoes National Park is now 100 times larger than before the invasion of alien grasses, even though an aggressive program of fire suppression by park staff has significantly diminished the sizes of individual fires. Rapid regeneration of grass overshadows the reestablishment of most native species, whose cover and abundance were drastically reduced for many years because of their sensitivity to fire. Leptecophylla tameiameiae (formerly called Styphelia tameiameiae), a common shrub in this ecosystem, is not fire resistant and is being extirpated from burned areas. Similar changes in fire cycles with grass invasions have occurred on the windward plain and Pūpūkea areas of O‘ahu and in the Puna and Ka‘ū regions of Hawai‘i. Schizachyrium condensatum (beard grass) is a summer-active grass that has become widely and densely established in dry forest areas of the Hawaiian Islands. Although not formally documented as present until 1961, it is thought to have been deliberately or perhaps accidently introduced in the 1930s or earlier. Schizachyrium now constitutes a major amount of standing dead biomass in dry forests of the island. Beginning in the late 1960s, the Hawaiian Islands experienced a major increase in both the number and size of fires. This surge in fire incidence and associated environmental damage can be directly attributed to the dry flash fuels of Schizachyrium. Because many native dry forest shrubs in the Hawaiian Islands have poor adaptations to fire, native species diversity is significantly lower after fire. The end result of invasions by alien grass is the conversion of the Hawaiian dry forest savannas to alien grasslands subject to frequent wildfires.

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A second species with a strong tie to fire is Andropogon virginicus (broomsedge), a perennial bunchgrass introduced more than 80 years ago from the southeastern United States. It readily invades open sites with a deep soil or ash deposit where the annual rainfall is above about 1,000 mm, where it can form continuous cover in a variety of habitats from dry lava fields to open mesic shrublands or even boggy areas. It continues to expand its range on all of the major islands today, from sea level up to 1,500 m elevation. The dead thatch produced each year by Andropogon is highly flammable, and its continuous cover provides an excellent fuel to carry fires over areas where they would not normally occur. This characteristic is evident where it has become established and carried fires into boggy areas of ‘ōhi‘a rain forest, despite the wet substrate. When not rejuvenated by fire, Andropogon individuals become senescent after 10–12 years of growth and die. Because Andropogon is a species that thrives on high temperatures, it grows during the hot summer months and goes dormant during the normally wetter winter months. The winter dormant period for Andropogon coincides with the wettest months in the Hawaiian Islands, and thus water is retained in the soil instead of being lost through evapotranspiration. Surface runoff results in increased rates of erosion, with slumping on steeper slopes where Andropogon has become densely established, as on windward O‘ahu. Under native evergreen rain forest canopies, water is transpired away rapidly and erosion and slumping are rare. Melinis minutiflora (molasses grass) is a spreading, mat-forming grass that gets its name from the sticky exudates secreted by glands over its leaf surfaces. It was originally introduced as a pasture grass but has now spread widely. Once established, Melinis forms pure stands that smother almost everything around. Experimental data have suggested that Melinis may be a superior competitor to Schizachyrium condensatum in summer-dry mesic forests, and that it will thus eventually replace much of the current biomass of this latter species. Melinis mats are adapted to fire and quickly resprout from their base and expand into adjacent areas. It is more fire prone than beard grass because it ignites under relatively moist conditions, due in part to

its glandular oils, increasing the likelihood that areas where it is established will burn. As with most grasses, its seeds are dispersed by wind. It is found on all islands from sea level to 1,500 m in dry to mesic environments, particularly in disturbed high-light areas.

grass with a densely clumped growth form and erect stems that grow to a meter tall. However, as in other parts of the world where it has been introduced, it quickly became a noxious weed, crowding out other herbs and seedlings where it has become established. C. setaceus produces huge quantities of seeds, which are dispersed long distances by wind. They readily colonize open sunny grasslands, dry forests, and lava fields from sea level to about 2,000 m elevation. This ability to colonize recent pahoehoe flows is not present in native grass species.

7.9  7.8 

Melinis minutiflora

Cenchrus clandestinus (kikuyu grass, formerly Pennisetum clandestinum) is an extremely aggressive perennial grass introduced to the Hawaiian Islands from Africa as a forage crop for cattle. Kikuyu grass reproduces both by seed and vegetatively using vigorous stolons and rhizomes. In the Hawaiian Islands, it quickly became naturalized in pastures, dry to mesic forests, and even disturbed open sites in wet forests at elevations up to 2,000 m on all of the major islands. Once established, its dense, smothering growth prevents virtually any new seedling establishment, and it has now become one of the most serious pest species threatening the restoration of native vegetation. However, because it forms dense mats, it burns poorly and generally retards fire. A second species of Cenchrus, C. setaceus (fountain grass, formerly Pennisetum setaceus), was introduced to the Hawaiian Islands from northern Africa as an ornamental. Its earliest documentation in the Hawaiian Islands was in 1914, but it may have been introduced as early as 1900. It is an attractive perennial

Cenchrus setaceus

Cenchrus setaceus alters ecosystem dynamics in three fundamental ways. First, its dense biomass aboveground makes it difficult for seedlings of native species to become established. Below ground, the dense root system of C. setaceus inhibits water and nutrient uptake by the roots of existing native species. Finally, and most significant of all, C. setaceus accumulates flammable fuel loads that increase the intensity and spread of fires and result in severe damage to native dry forest species. As with other fire-adapted grasses, it quickly resprouts after fire and reestablishes dense growth. Today, much of the dry leeward side of Hawai‘i is covered by a dense monoculture of Cenchrus, with major negative impacts on the remaining dry forest elements in this area. There have been extensive efforts to restore the natural ecosystem structure of dry and open mesic forests, particularly in Volcanoes National Park and on the slopes of Mauna Kea. Restoration efforts have had only limited success due to the delicate balance between keeping some level of nonnative herbivores to control the density of grass but not so many that they consume all koa and other seedlings.

A L I E N I N VA D E R S

Mesic Seasonal Forest and Wet forest Mesic and wet forests in the Hawaiian Islands have received less human disturbance than dry forests and open seasonal forests. The Native Hawaiians viewed the upper-elevation rain forests as the realm of the gods and thus impacted these areas only through limited hunting and gathering activities. The arrival of Europeans put more pressure on these forests, particularly with the sandalwood trade in the early nineteenth century. By the end of the nineteenth century, sugar growers in the Hawaiian Islands realized that their industry depended on open irrigation ditches to bring water from the upper forest slopes to the cane fields below. With the annexation of Hawai‘i by the United States in 1902, the territorial government established the Division of Forestry to create forest reserves for the primary function of protecting water resources, with wood production given secondary importance. Harold Lyon, hired as a plant pathologist by the Hawaiian Sugar Planters’ Association (HSPA) in 1907, came to play a major role in the introduction of alien tree species to these reserves, as described in chapter 8. Lyon felt that native trees were poorly adapted to changing forest conditions and disturbance and that their decline was inevitable and irreversible. Thus, he felt the protection of these watersheds required planting alien trees. Over the course of nearly half a century, HSPA cooperated with the Division of Forestry to introduce large numbers of tree and shrub species to the Hawaiian Islands. In the end, the species planted most frequently were those that were easy to establish, grew rapidly, and seeded abundantly. Among species introduced to forest reserves, a number have proved to be highly invasive into native forests. Morella faya (fire tree, Myricaceae), formerly named Myrica faya, presents a classic example of how an invasive alien species can fundamentally alter the structure and function of a native forest ecosystem. This shrubby tree can grow to as much as 12 m in height and is native to the Canary Islands, the Azores, and Madeira, where it grows on volcanic soils that resemble those of the Hawaiian Islands. A major basis for its original introduction in the early 1900s by local

Portuguese was its use as an ornamental and for the production of wine. However, a key trait of M. faya is that it possesses root nodules that host bacteria called actinomycetes. These bacteria fix nitrogen directly from the air, allowing a ready source of nitrogen to the host plant and thus a means of colonizing nutrient-poor soils. With these traits, Morella was considered a perfect choice for reforestation of cutover forestlands and open forest areas. However, it proved so successful on such sites that it has become an aggressive invader of open forest areas. Morella fruit is dispersed by alien and native fruit-eating birds as well as by feral pigs, with the seed remaining viable in the soil for a long period. Not only does Morella choke out native shrubs and tree saplings, it enriches the otherwise poor soils and thereby provides conditions that facilitate the establishment of other invasive species, particularly grasses. From the 1960s through the 1980s, Morella expanded explosively into pioneer Metrosideros forests on young lava flows on Hawai‘i, where it quickly became a codominant species. Some dieback from insect pests has since occurred, and the future role of Morella in these forests remains unclear.

7.10 

Morella faya

Another species with the ability to fix nitrogen through symbiotic root nodules is Falcataria moluccana (Moluccan albizia; Fabaceae; formerly known as Albizia moluccana). This tree species, which grows rapidly to up to 40 m in height, was introduced to Hawai‘i from Indonesia by Joseph Rock in 1917 as an ornamental tree and for reforestation. Extensive

plantations were established in mesic to wet lowland areas of Kaua‘i and secondarily on O‘ahu and Hawai‘i. From these plantations, it has spread slowly into mesic forests, pastures, and disturbed areas. Its impact on native tree species comes with its rapid growth rate and broad canopy that shades out slow-growing species such as Metrosideros polymorpha. Psidium cattleianum (strawberry guava; Myrtaceae) was introduced as an ornamental in 1825 and soon became established in the wild. On most islands, nearly pure thickets of this species infest hundreds of hectares of mesic and welldrained rain forest areas at elevations of 200–1,300 m. Vast areas of mature ‘ōhi‘a and koa forests today have a dense understory of P. cattleianum, making this the worst alien invader of wet forests. Once P. cattleianum is established, few native species can survive in the dense shade under its thickets. Moreover, feral pigs relish eating the guava fruit during the fruiting season, thoroughly disturbing the ground and trampling any surviving understory plants. Guava seeds then pass through the digestive system of the pigs and are dispersed to new sites. Alien fruit-eating birds also carry the seeds to new areas. A related species of guava, Psidium guajava, colonizes disturbed areas of wet to moderately dry forests at low elevations on all of the major islands, where it forms dense thickets much like those of strawberry guava.

7.11 

Psidium cattleianum

The presence of Schinus terebinthifolius (Christmas berry; Anacardiaceae), a South American tree widely invasive in subtropical areas throughout the world, was first noted in the Hawaiian Islands in 1909. It was likely introduced as an ornamental tree. The red berries that give this species its name are widely dispersed by alien birds, with a resulting rapid increase

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in its range in wet and mesic lowland environments. Once established, it forms dense stands up to 8 m tall that shade and choke out other species. Its appearance as a community-dominant tree is relatively recent in the Hawaiian Islands, where it seems to outcompete even other invasive alien trees. It has been reported to hold its own in competition even with Psidium guajava, and it replaces Leucaena at its wetter margin of occurrence. Schinus is killed by high-intensity fires but regenerates rapidly where there is a large seed bank. Schefflera actinophylla (octopus tree; Araliaceae) is a fast-growing, multistemmed tree with few sparse branches and distinctive palmately divided leaves. This tree, which reaches heights of 15 m, was introduced to the Hawaiian Islands a century ago and is widely cultivated on all of the major islands. The seeds are dispersed by alien fruit-eating birds, which carry them to wet forest areas at low elevations where the shade-tolerant plants are capable of invading undisturbed forests. Schefflera can be found growing in wet lowland habitats up to 1,000 m elevation on the larger Islands. There are major infestations in the northern valleys of Kaua‘i, particularly Limahuli Valley, and in Nu‘uanu and Waiāhole valleys on O‘ahu. These trees have the interesting capacity to grow as epiphytes in wet forests, with seedlings readily colonizing crotches of tree branches and moist cliffs. Schefflera is well known as a serious pest in Florida, where it aggressively invades intact, undisturbed hardwood forests and, to a lesser extent, pine woods.

Ficus microcarpa (Chinese banyan; Moraceae) is widely planted as a park or shade tree throughout the world, with the Hawaiian Islands as no exception. It was first introduced early in the twentieth century but became naturalized only after about 1938, when the wasp species that serves as its specific pollinator was also introduced. The presence of these wasps allowed F. microcarpa to produce viable seeds and spread from initial plantings. Harold Lyon favored this species strongly in his plantings, and it is one of the few figs to become an aggressive invader. Typically, F. microcarpa grows as a strangler fig, with the seedlings beginning their lives as epiphytes on a host tree. Eventually the plants’ aerial roots reach the ground, and newly rapid rates of growth provide tissues that slowly engulf their host. Ficus microcarpa is capable of growing in a variety of conditions from wet to dry in disturbed, low elevation habitats. As urban areas are built closer to natural areas, F. microcarpa will increase as an invader of native forests. It does not invade undisturbed forest, but once established it will displace all other trees with its dense shade. The fruit of this species is small and is eaten by numerous fruit-eating birds and other animals.

7.13 

7.12 

Schefflera actinophylla

Ficus microcarpa

Spathodea campanulata (African tulip tree; Bignoniaceae) was introduced to the Hawaiian Islands as an ornamental tree in the middle of the nineteenth century. It is so widely planted as a roadside and shade tree today that many assume it is a native species. The environmental problems presented by this species are not caused by these urban plantings but by the winged seeds that float great distances on the wind. Seedlings of Spathodea readily become established in native

forests along stream courses, where they thrive because of their ability to tolerate shady understory conditions. Adult trees with their broad canopies shade out native species. Today, African tulip trees are widely established on most of the major islands in moist to wet low-elevation forests, with major infestations tucked away in almost every rain forest valley along the northern and eastern slopes of Kaua‘i, O‘ahu, and East Maui. Increasing numbers are evident in windward Hawai‘i valleys as well.

7.14 

Spathodea campanulata

Corynocarpus laevigatus (karaka, New Zealand laurel; Corynocarpaceae) is another aggressive tree invader in mesic forests, introduced because of its ability to colonize open sites. These trees readily colonize open areas and grow to 15 m in height and diameters up to 60 cm, with a dense evergreen canopy that shades out native species. It was actively planted on Kaua‘i in the late nineteenth and early twentieth centuries, with reports even of aerial seeding. Today it is a major threat to native forests on Kaua‘i, but it is also present in smaller numbers on O‘ahu, Moloka‘i, and Hawai‘i. The fleshy fruit is edible, and this leads to its spread by pigs and frugivorous birds. The seeds, how­ ever, are highly toxic. The Melastomataceae, a family filled with many plants specialized for the colonization of disturbed habitats, includes some of the most invasive and potentially damaging species of invasive aliens in the Hawaiian Islands. Of the 15 melastome species naturalized in the Hawaiian Islands, 9 have been declared noxious weeds. The worst of these are Clidemia hirta and Miconia calvescens. Several other melastomes that are widespread in cultivation have been declared as noxious weeds and deserve careful attention.

A L I E N I N VA D E R S

These include Tibouchina herbacea, T. urvilleana, and Oxyspora paniculata. As its common name Koster’s curse indicates, Clidemia hirta represents a major environmental threat to wet forests in the Hawaiian Islands. It is not known for certain when or how this melastome reached the Hawaiian Islands, but it was first reported as present in 1941. This weedy shrub grows up to 2 m tall in dense thickets on open areas and semishaded forest habitats, especially along trails and roads. It is an aggressive invader that shades out all plants below it, including mosses, lichens, and low herbs that act to bind forest soils and protect them from erosion. The seeds are principally dispersed by alien fruit-eating birds but also by mammals such as pigs and humans, who carry seeds as they move along trails. Each fleshy fruit contains more than 100 tiny seeds that can easily become attached with mud to shoes and car tires. There is evidence that marijuana growers inadvertently establish many new infestations. Although not resistant to fire, it rapidly colonizes burned areas. Clidemia hirta is a serious pest in mesic and wet environments on O‘ahu and more recently in Wailau Valley on Moloka‘i, as well as the Nāhiku and Kailua areas on Maui. It has also become established on Kaua‘i, West Maui, and Waiākea, Hawai‘i.

7.15 

calvescens is a small tree up to 15 m tall, with leaves that can reach more than 2 m in length. The canopy created by these huge leaves prevents light from reaching the forest floor, effectively excluding all other plants. It has become an aggressive invader in local areas of moist to wet forests on Kaua‘i, O‘ahu, Maui, and Hawai‘i. Once established it produces copious amounts of fruit with small seeds, spread primarily by alien fruit-eating birds. Although M. calvescens readily becomes established on steep slopes of wet forest areas, its root system is shallow and prone to sliding in heavy rains, and thus it contributes to landslides and major erosion. The potential impact of this alien tree on native biodiversity in the Hawaiian Islands can be seen in Tahiti, where M. calvescens was introduced in 1937. There it rapidly invaded native forests to the extent that two-thirds of these forests today are dominated by M. calvescens, with disastrous consequences for the native flora. Perhaps no other single invasive plant species has so great a potential for serious environmental damage to wet forests of the Hawaiian Islands.

Clidemia hirta

A second member of the Melastomataceae, which has been the cause of great environmental concern in recent years, is Miconia calvescens (velvet tree). It was introduced innocently to the Hawaiian Islands from tropical America as an ornamental tree in 1961, but by the early 1990s it had escaped cultivation and become widely naturalized. Miconia

7.16 

Miconia calvescens

Several species of blackberry have become serious invaders in the Hawaiian Islands, with 2 of these causing major changes in mesic and wet forest eco­

systems. Rubus argutus (Florida prickly blackberry; Rosaceae) is a thorny scrambler that rapidly invades disturbed areas and has become well established in mesic to wet forests from 200–2,300 m on five of the eight major islands. Once it colonizes an area, its aerial shoots readily root to form impenetrable thickets. Alien fruit-eating birds are the principal dispersal agents. The aboveground tissues of the plant are destroyed by fire, but the plant quickly recovers from basal and subterranean shoots. Another species of recent concern is Rubus ellipticus (Himalayan raspberry; Rosaceae), a trailing bramble that was introduced to Volcano, Hawai‘i, in 1961 from Asia. Today it is well established in that region of the Big Island in wet forests from 700–1,700 m elevation, principally in pig-disturbed areas. Rubus ellipticus spreads by vigorous vegetative growth as well as by seeds dispersed by both alien and native birds and mammals. It is hard to kill once established and forms impenetrable thickets. These threaten native wet forests and displace native plant species, including the native Hawaiian raspberry, Rubus hawaiiensis. Although not yet naturalized in similar forests on Maui, it would likely be a serious problem if it becomes established. Not all aggressive invaders in moist and wet Hawaiian forests are trees or shrubs. One notable species of major concern is the vine Passiflora tarminiana (banana poka; Passifloraceae; formerly called P. mollissima). Initially introduced to the Hawaiian Islands from South America early in the twentieth century for its ornamental flowers and edible fruit, P. tarminiana has become one of the worst forest invasives, covering thousands of hectares of moist forest on Kaua‘i and Hawai‘i. On Maui, P. tarminiana is currently restricted to a few square miles in the Kula Forest Reserve area, but it is expanding its range. The fleshy fruit is widely sought after by pigs and alien fruit-eating birds, which disperse them widely. Once established, Passiflora tarminiana overgrows native koa and ‘ōhi‘a, smothering them under its foliage. Where the native tree canopies have been opened, dense mats of Passi­ flora vines cover the understory trees and shrubs and inhibit regeneration of native trees. As many as 30 species of Passiflora have been introduced in the Hawaiian

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Islands, and 13 of these have become naturalized. A second species of particular concern is Passiflora suberosa (huehue haole), which like P. tarminiana is dispersed by alien fruit-eating birds. It can grow to smother both upper and lower canopy layers in dry forests and occurs in dryland habitats on all the major islands between sea level and 600 m, with major infestations throughout the Wai‘anae Mountains on O‘ahu and along the Kahoma Ditch trail on Maui.

Naturalized gingers are widespread today in wet habitats from sea level to 1,700 m. There are major infestations at Kōke‘e on Kaua‘i, Nāhiku on Maui, and Volcano on the Big Island. Two other Hedychium species, H. coronarium (white ginger) and H. flavescens (yellow ginger) are widely naturalized and thus species of concern as well.

7.19 

Sphaeropteris cooperi

A second large alien fern species of concern is Angiopteris evecta (mule’s-foot fern; Marattiaceae), which has a stemlike rhizome up to 1 m in height, with leaves to 3 m or more in length and 2 m in width. Angiopteris has been widely cultivated in the past and is now freely naturalizing in wet valleys in the Ko‘olau Mountains on O‘ahu, spreading by spores.

7.18 

7.17 

Passiflora tarminiana

Several species of gingers, most notably Hedychium gardnerianum (kāhili ginger; Zingiberaceae), have become aggressive weeds in the Hawaiian Islands. This showy ginger, which grows to about 1 m tall, is widely planted in tropical gardens throughout the world. The conspicuous, fleshy, red seeds are dispersed by alien and perhaps native fruit-eating birds. Inadvertent human dispersal also occurs, as even small discarded rhizome fragments will root and form new plants. Hedychium prefers mesic open-light environments, particularly along stream channels, but it will readily grow in semiand full shade beneath forest canopies. Once established, stands of Hedychium gardnerianum expand rapidly through vegetative propagation, forming dense thickets that displace all other plants.

Hedychium gardnerianum

Ferns and fern relatives are an important component of the native forests of the Hawaiian Islands. Many more ferns have been cultivated over the past two centuries, and at least 37 of these are reported to have become naturalized. One of these ferns that has become a serious invader is Sphaeropteris cooperi (Australian tree fern; Cyatheaceae). This large species, which can reach 9 m or more tall, was introduced to Hawaiian gardens more than 60 years ago and continues to be a standard species in the nursery trade. Unlike many invaders, S. cooperi can readily become established and grow rapidly in undisturbed native wet forests, as on the windward upper-slope forests of Kaua‘i that are otherwise relatively free of alien species. Because its windblown spores may disperse great distances, it can readily become established in areas away from human or pig disturbance.

Upper Montane, Subalpine, and Alpine Ecosystems Human activities and introduced grazing animals have had major environmental impacts on higher elevation areas in the Hawaiian Islands since the middle of the nineteenth century, when large herds of cattle and goats grazed the upper slopes of Mauna Kea and Mauna Loa. Although the impacts of invasive plant species on native ecosystems have not been as great in the lower-elevation dry and wet forests, more than 150 naturalized plant species have been recorded at elevations above 2,000 m on Maui and Hawai‘i. Not surprisingly, these naturalized upper-­ elevation species have origins largely from temperate areas of the world. Several of the potentially disruptive invaders are temperate forest trees that have escaped upward from plantations, as with Pinus radiata (Monterey pine; Pinaceae) and Eucalyptus globulus (blue gum; Myrtaceae).

A L I E N I N VA D E R S

An alien shrub species that, like Morella, has the ability to fix atmospheric nitrogen and fundamentally alter soil nutrient conditions is Ulex europaeus (gorse; Fabaceae). The spiny legume shrub native to Western Europe and the Mediterranean Basin has become widely established in higher elevation grasslands and degraded shrublands on both Maui and Hawai‘i. If left to expand, this species can form virtually impenetrable thickets. It readily responds to fire by resprouting from its base and by seedling germination from soil seed pools. A widespread invasive grass at higher elevations is Holcus lanatus (common velvet grass; Poaceae), a perennial bunchgrass that readily invades disturbed sites. This species is native to northern Europe. Seeds are produced abundantly and dispersed by wind, with subsequent rapid growth of seedlings that form dense stands and outcompete native species. As with most bunchgrasses, this species tolerates fires and regenerates rapidly from basal shoots. Holcus is widely distributed in all but the driest montane habitats, most commonly above 1,300 m. There are indications that Holcus is now a permanent component of Hawaiian montane grassland communities and will remain even if disturbance is eliminated. It also readily colonizes montane bogs, particularly following damage by pigs. Verbascum thapsus (woolly mullein; Scrophulariaceae) is a biennial rosette plant with woolly foliage, native to central and northern Europe. It becomes invasive on the Hawaiian Islands by quickly colonizing disturbed areas with its wind-­

dispersed seeds, forming a dense ground cover that displaces slow-growing native species. Verbascum infests roadsides at elevations from 1,500 to 3,000 m on Mauna Loa, Mauna Kea, and Hualālai and is particularly dense around 2,000 m, where it can form a cover that can out compete native vegetation. The massive seed production and dispersal make its eradication extremely difficult. Careful monitoring has kept this species from colonizing similar habitats on Haleakalā, where it would likely become a major invader were it to become established.

Shoreline Mangroves Although mangroves are widely present on Pacific Islands, no species were able to reach the Hawaiian Islands through natural dispersal. Rhizophora mangle (red mangrove; Rhizophoraceae) was introduced to the Hawaiian Islands in 1902 to promote shoreline stabilization of mudflats on southwestern Moloka‘i. From these original introductions, seedlings were able to disperse widely to most of the other major islands, and extensive stands have developed on O‘ahu. In 1922, Brugiera sexangula (oriental mangrove, Rhizophoraceae) was introduced to He‘eia on O‘ahu, where it continues to persist. Rhizophora has thrived in shallow inlet waters of the Hawaiian Islands that are highly disturbed by human actions and offer few competitors or herbivores. As intended with their original introduction, areas where mangroves grow and dominate slow the loss of sediment and organic material from the shore,

changing the fundamental structure of these ecosystems. These stands have significantly altered brackish water ecosystems as well as many fishponds and have had dramatic effects on communities of intertidal invertebrate and native shorebirds. Mangroves are now being controlled because of their interference with shoreline archaeological sites and disruption of the habitat of endangered bird species.

The Ten Worst Invaders Considering the species of invasive alien plants described above, which have the greatest negative impact on native Hawaiian ecosystems today? There might be some differences of opinion about the precise composition of such a list. A first approximation of this might include the following, in alphabetical order: Andropogon virginicus, Clidemia hirta, Leucaena leucocephala, Miconia calvescens, Morella faya, Passiflora tarminiana, Cenchrus clandestinus, C. setaceus, Psidium cattleianum, and Schinus terebinthifolius. However, many would argue that Rubus argutus, Schizachyrium condensatum, and Sphaeropteris cooperi would also be worthy candidates to make this list. Nevertheless, invasives are a dynamic group of species, and this list will surely change in the future. Effective biological control agents have reduced the impacts of some species, as with Lantana camara, which would have once made this list, while other species such as Melinis minutiflora may well move onto the list in the future.

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Limahuli Valley, an ahupua‘a on Kaua‘i

CHAPTER 8

Conservation of the Hawaiian Flora speciation remained slow in evolutionary time. Our best evidence is that perhaps hundreds of unique plant species and at least half of the native birdlife fauna have been lost since the first human arrival on the Islands. There have undoubtedly been thousands of lesser-known taxa of insects, spiders, mollusks, and lower plants that were lost before they were ever described. Moreover, many species today exist in small numbers in relatively few populations and thus are easily threatened by extinction. Today, with less than 0.2% of the land area of the United States, the Hawaiian Islands hold more than a third of the nation’s entire listing of endangered and threatened species under the U.S. Endangered Species Act.

Hawai‘i’s Unique Biota The Hawaiian Islands, with their location in the middle of the Pacific Ocean more than 3,700 km from the nearest continent, are arguably the most isolated archipelago in the world. Those organisms that were able to reach the Islands and become successfully established found a landmass with an exceptional diversity of climatic and geological features that provided an enormous range of available habitat conditions. The slow colonization of the Hawaiian Islands by long distance dispersal could have begun as early as about 28 Ma with the formation of Kure Island, but only low and relatively short-lived islands existed until the emergence of Lisianski Island about 23 Ma. However, the majority of early colonizers would have been lost as the older islands eroded and high-elevation landmasses were eliminated. It was only with formation of Kaua‘i about 4.7 Ma that a stable platform for colonization of our modern lineages was present. For those species successful in dispersing to the Hawaiian Islands and becoming established, gene flow from their distant parent populations would have been low to even nonexistent, while at the same time the new environments would have provided strong selective pressures for adaptation. It is for this reason that the Hawaiian Islands provide such classic textbook examples of speciation and adaptive radiation. Although Charles Darwin was aware of the unique biology of the Hawaiian Islands, he never visited the Islands during his voyage on HMS Beagle. Had he encountered the Islands, he would have discovered Hawai‘i to be arguably the world’s premier showcase of adaptive radiation, surpassing the Galapagos Islands in the number and variety of species that have evolved from a small set of colonizing ancestors. Levels of endemism of about 90% for vascular plant species are unmatched for any other major area in the world. High levels of endemism are also present in terrestrial invertebrates and bird species, and even marine species of the Hawaiian Islands exhibit endemism levels of 15–20%, a remarkably high level for islands. We know that almost certainly there have been thousands of Hawaiian species that evolved and later became extinct over the long geologic history of the Islands. However, the size of the modern flora and fauna is a clear indication that rates of speciation have exceeded rates of extinction over this period. Nevertheless, with the arrival of man in the Islands and associated changes in land use and habitat degradation, these patterns were reversed and rates of extinction accelerated quickly in ecological time, while the rate of

An Endangered Flora The evolutionary history and geographic isolation of the Hawaiian Islands have predisposed the flora to characteristics of distribution, population structure, and reproduction that have inherently produced fragile conditions for species survival. This aspect of isolation can be readily seen, as it applies to geographic range and habitat occurrence. Among endemic flowering plant species in the Hawaiian Islands, 68% have a range restricted to a single island, and 78% of species exhibit relatively narrow ranges of habitat distribution. Population numbers and population size represent another area of sensitivity. Remarkably, or perhaps a better term would be alarmingly, nearly 25% of the Hawaiian species of vascular plants (including species that have become extinct in historical times) are known from fewer than 100 individuals. This rarity can be seen in Delissea argutidentata, which once reached heights of 10 m and grew on the slopes both of Mauna Loa and Hualālai. Today it is known from a single individual. More than 40% of the species are known from less than 1,000 individuals. This rarity applies to populations as well, with 40% of the species known from only 10 natural populations or less. There are 91 species known from only a single population, putting their prospects for survival at extreme risk. If one analyzes the species categorized as at risk in the Hawaiian Islands today, as discussed in this chapter, species on this list are significantly likely to share one or more of the following three components of rarity. These are limited geographic distribution, highly specific habitat requirements, and small population size. Compared to secure taxa with stable populations, species at risk are commonly single-island endemics, restricted in distribution to specific habitat types, and occurring in small population sizes and numbers.

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8.1  Delissea argutidentata, photo from Joseph Rock’s 1919 monograph on the Hawaiian lobeliads

Small, isolated populations are well known to potentially experience demographic problems of reduced levels of genetic variability, which diminish the capacity of a species to adapt and respond to disturbance or environmental changes and thereby lowers the probability of long-term persistence due to genetic bottlenecks. Species with small and isolated plant populations may also experience reduced reproductive vigor due to inbreeding depression or ineffective pollination. These genetic constraints of small population size are notable for dioecious species where male and female flowers occur on separate individuals, although dioecy has the benefit of ensuring outcrossing. The problems of Hawaiian species that are associated with small population size and vulnerability to random demographic fluctuations or natural catastrophes are further magnified by synergistic interactions with other threats. Thus, many species with local geographic ranges are inherently vulnerable to extinction because of disturbance, climate change, feral grazing animals, random demographic fluctuations, hurricanes, and disease outbreak.

In contrast to species considered at risk, opposite traits and patterns of occurrence are associated with species that appear to have stable populations and that are relatively secure in their survival. Acacia koa and Metrosideros polymorpha are good examples of secure species with multi-island patterns of distribution, wide-ranging habitats of occurrence, and large population numbers and sizes. Most indigenous native species also fit this category of secure status because of related traits, with 95% of these indigenous Hawaiian species having multi-island distributions and most occurring in relatively large population numbers and sizes. As a result, only a small number of indigenous species (less than 4%) are considered to be at risk by definitions described here. While geographic ranges of occurrence on multiple islands and broad ecological distribution are generally associated with species more secure in their survival, there are exceptions to this generalization. Good examples of these exceptions can be seen in a number of dry forest trees that were once widespread with moderate to large populations on most of the major islands. Here, human history and land use have not been kind to this ecosystem. Historical factors of habitat destruction and the loss of native bird species associated with pollination or seed dispersal have pushed some species to a threatened status. Many endemic Hawaiian bird species that are now extinct coevolved with particular plant species, with bills of just the right length and curvature to pollinate them. This is particularly easy to see among the lobeliads, whose flowers are mimics of those bird bills, but there are many other examples as well, including native species in the hibiscus family for which no native pollinators have survived. Additionally, many insect pollinators are now extinct, potentially leaving their plants without the proper vector to transfer pollen between the flowers. Many organizations and agencies have been established to help preserve rare species. The establishment of the National Pacific Botanical Garden on Kaua‘i in 1970, under a charter from the U.S. Congress, has had a major impact on the conservation of rare species in the Hawaiian Islands and the broader Pacific region. Also important is the Plant Extinction Prevention Program (see pepphi.org), which works through collaboration with large and small private, state, and federal landowners, land managers, researchers, and other partners. A consortium of over 35 conservation agencies and organizations and private landowners provide oversight for the program. Many species survive today because of these programs to protect rare plants. Nevertheless, the long-term future of these species is in doubt, as many survive only in cultivation or by extreme measures of hand pollination in the field.

Categories of Rarity In working to promote the preservation of native Hawaiian plant species, it is useful to categorize species that we would designate as being at risk, in contrast to those whose survival seems relatively secure. Several assessments of this type have been made (see appendix). The risk assessments used on the Smithsonian Web site and adopted in the text were developed in a collaboration of the U.S. Fish and Wildlife Service, the Nature Conservancy, and Hawaiian field botanists (Wagner et al. 1999). Additional

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information can be found in the Plant Red Data Book produced internationally by the International Union for the Conservation of Nature (IUCN) and regularly updated for the Hawaiian flora by the Natural Heritage Program of the Nature Conservancy of Hawai‘i. The more formal federal categorization of endangered and threatened species is established by the U.S. Fish and Wildlife Service under the Endangered Species Act. Although the categories used in these two systems differ slightly, their overall structure for the delineation of at risk species is quite similar. However, federal listings are fewer because adding species under the Endangered Species Act is very slow and influenced at times by political pressures. We have adopted four categories of conservation threat for native species considered to be at risk using those delineated by Wagner et al. (1999) with appropriate updates (see appendix). The most extreme level of risk is that of extinct, a category for those taxa that are thought to no longer occur in the wild. There are 118 Hawaiian plant taxa in this category, or about 9% of the total historically known vascular plant taxa. A notable example of species now extinct is Cyanea arborea, a tall palmlike species that once occurred on the leeward slopes of Haleakalā. To provide some perspective on the significance of this figure, less than 1% of the known California flora is considered to be extinct in the wild. In a few cases, these Hawaiian species that no longer exist in the wild have been propagated in the past and survive in cultivation. Examples of these include Silene pearlmanii, Hibiscadelphus giffardianus and H. hualalaiensis, and Kokia cookei. Plant conservation in botanical gardens and other living collections is referred to as ex situ conservation as opposed to in situ conservation that occurs in natural habitats. The second category is that of endangered species, being those taxa in danger of becoming extinct throughout all or a significant part of their range in the Hawaiian Islands unless threats to their survival can be alleviated. The group includes about 315 taxa, or nearly one-quarter of the flora. An extreme example is the rare endemic Kanaloa kahoolawensis known from just 2 mature living individuals, one on Kaho‘olawe and one in a botanical garden, although Kanaloa pollen shows up in cores from Kaua‘i, O‘ahu, and Maui. Similarly illustrating the unpredictable rarity of some species, Lysimachia venosa was originally known from just 2 collections from the summit of Mount Wai‘ale‘ale, Kaua‘i, the last in 1911. Eighty years later in 1991, a broken branch was discovered at the base of 1,000 cliffs at the headwaters of the Wailua River, Kauai. The branch was from L. venosa and was assumed to have fallen from the summit of Mount Wai‘ale‘ale. No more specimens were known until the discovery in 2012 of about 30 plants growing on the windward slopes of Kaua‘i just below the summit of Kawaikini. Vulnerable species are defined as those that are likely to become endangered unless the circumstances threatening their survival and reproduction improve. This group comprises about 60 taxa or 4.4% of the flora. Finally, rare taxa form a group of species with small and localized populations or highly specific habitats that are not presently considered to be endangered or vulnerable but are considered as potentially at risk because of these restricted habitat ranges and population sizes. There are about 163 taxa in this category, or about 12% of the flora.

Combining the four categories, nearly half of the taxa in the flora are in categories classified as at risk. Even at the species level, more than 40% of the flora is at risk. Several genera provide examples of concern about rarity and extinction. For example, 11 of the 15 known species of Delissea (Campanulaceae) are considered to be extinct in the wild today, while the other 4 species are all classified as endangered. A similar pattern of species loss can be seen in smaller genera such as Hibiscadelphus (Malvaceae), with 6 of 7 species thought to be extinct in the wild and 1 endangered, and Haplostachys (Lamiaceae), with 4 of the 5 known species extinct and the fifth endangered. Even large genera can show extreme patterns of rarity. Of the 34 species in Schiedea (Caryophyllaceae), 2 are extinct today but 31 are classified as endangered, rare, or vulnerable. Only a single species is considered reasonably secure in its populations. At the federal level, there are two categories of listing that have official status under the U.S. Endangered Species Act and are thus protected by statutes that require conservation plans with designations of critical habitat to help protect these species. The first of these federal categories is that of endangered species, defined by law as those in danger of becoming extinct throughout all or a substantial portion of their range unless the threats jeopardizing their survival are mitigated or eliminated. With recent designations by the U.S. Fish and Wildlife Service, 362 taxa, or about 27% of the Hawaiian flora, is categorized as endangered. Threatened species, roughly equivalent to vulnerable species in our use here, forms the next category under the Endangered Species Act. These are defined as species likely to move into the category of being endangered in the near future unless threats to their survival are reduced. There are 12 taxa in this category, or about 1% of the flora. There is no federal category for rare species. Species or taxa in the process of consideration for protected federal status are termed “candidate species,” and there are 38 taxa in this category. Another approach to listing plant species of conservation concern has been carried out by the State of Hawai‘i under a program called the Comprehensive Wildlife Conservation Strategy. Species listed were chosen as Species of Greatest Conservation Need (SGCN) based on the following criteria: (1) plant species federally listed as threatened, endangered, or as a candidate for listing; (2) plant species identified as Genetic Safety Net (GSN) plants (i.e., plants with less than 50 individuals); (3) plant species identified as important elements of native habitats; and (4) endemic aquatic plants. This analysis listed about 550 taxa as SGCN and 177 as GSN plants because of their small population size.

Threats from Habitat Destruction and Land Cover Changes Although there exist threats to specific species, the major concern with the preservation of the native Hawaiian flora overall relates to issues of habitat destruction. Native Hawaiian ecosystems have undergone dramatic and severe changes in land use and habitat destruction that began with the arrival of the Polynesians and have continued through the period of Euro-American colonization up to the present time. Evidence obtained both from archaeological studies and paleoecological research with pollen cores suggests that the prehistoric Polynesians had a far greater impact on the Hawaiian

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8.2 

Nothocestrum breviflorum

ecosystem than had once been realized. Polynesian colonization of the Hawaiian Islands began with the introduction of alien plants and animals and quickly moved to changes in land cover as agriculture developed. As Hawaiian populations grew rapidly, most land with even moderate soil fertility below about 600 m elevation was cultivated between the thirteenth and nineteenth centuries. The cumulative effects of this forest clearance and habitat modification through the use of fire led to major changes in lowland ecology, although there is little or no evidence of major impacts to vegetation at higher elevations. Some examples of vegetation and ecological changes resulting from the activities of the Polynesians are the decline and alteration of the structure of lowland dry forests, increased erosion, lowland plant extinctions and extirpations, loss of many endemic bird species, destruction of once-extensive Pritchardia palm forests, and the near-extinction of Kanaloa kahoolawensis. The later arrival of Euro-American colonization led to the intensification of agriculture and resource extraction through the commercial exploitation of sandalwood and firewood for whaling ships, the introduction of goats and European pigs, and the development of cattle ranching. As agricultural production

8.3 

Colubrina oppositifolia

of sugarcane and pineapple increased, efforts to protect watersheds for commercial irrigation led to the widespread planting of nonnative tree species beginning in the early 1900s. Many of these introduced tree species are invasive in Hawaiian forests today (see chapter 7). No forest community in the Hawaiian Islands has undergone greater change and degradation due to human activities than the dry forest communities. Although dry forests once occurred on the leeward aspect or rain-shadow side of the main islands and were once highly diverse in species composition, changes in land use and land cover, grazing by nonnative ungulates, establishment of invasive alien plant species, and fire have combined to cause severe degradation and fragmentation of remaining stands. Many dry forest trees that were once common are highly restricted in occurrence and rare today, as with the endangered Nothocestrum breviflorum on Hawai‘i and Colubrina oppositifolia on O‘ahu, West Maui, and leeward Hawai‘i. The largest remaining stands of dry forest today occur on Kaua‘i and Hawai‘i, but they are highly disturbed and cover less than 2% of the land area. O‘ahu, Lāna‘i, and Maui contain only small degraded fragments of their once extensive dry forest communities.

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One of the greatest impacts to Hawaiian ecosystems has come from introduced nonnative mammals. The first introductions of alien mammals began with pigs, dogs, and rats that arrived with the Polynesians, and later came the introduction of larger grazing ungulates following the arrival of Captain James Cook in 1778. Early explorers and later colonists intentionally introduced European pigs and other livestock such as goats and blacktailed deer, mouflon sheep, and cattle. Up until relatively recent actions to reduce their effects, these feral ungulates dramatically increased in numbers and distribution across the Islands, with major impacts on native plant communities. Beyond the direct ecological effects of trampling and consuming native vegetation, the actions of feral ungulates have contributed significantly to increased erosion and facilitated the spread and establishment of invasive alien plant species. It was reported that at one time, ranchers on Maui deliberately overgrazed the land so that soil would erode and slide down the hill, thus creating a wider area for pasture at low elevations; this was also done at Mānā, Kaua‘i, for sugarcane planting. The European pig, first introduced to the Hawaiian Islands by Captain Cook, quickly hybridized with domesticated Polynesian pigs and became feral and well established in many wet and mesic forests, as well as drier areas at high elevations. Feral pigs have arguably had the most pervasive and disruptive nonnative influence on the native forests of the Hawaiian Islands, and they continue to be a significant threat to many mesic and wet forest areas despite active hunting programs. Pigs have also impacted Hawaiian bogs, where they are apparently attracted by the availability of alien earthworms, which provide a ready source of protein. The removal of plant cover in bogs by pigs can approach 100%. Pigs directly impact native plants by rooting for fleshy tissues and invertebrates, disturbing and destroying vegetative cover, and trampling plants and seedlings. Pigs also inhibit or

prevent native plant regeneration by damaging or eating seeds and seedlings. Moreover, pigs often selectively seek out specific plant species, and those with fragile stems and leaves have been dramatically reduced in numbers in many areas. Tree ferns and other ground ferns have been particularly impacted. In addition to their direct effects on vegetation, pigs are a major vector for the establishment and spread of invasive alien plant species. They disperse seeds of alien plants on their hooves and coats as well as internally through the spread of manure and by fertilizing the disturbed soil through their feces. Passiflora tarminiana (banana poka) and Psidium cattleianum (strawberry guava), and Clidemia hirta (Koster’s curse) are examples of alien species widely spread by pigs. Feral pigs are currently present on Kaua‘i, Ni‘ihau, O‘ahu, Moloka‘i, Maui, and Hawai‘i, although their numbers have been significantly reduced from former levels within fenced areas. Goats were first introduced to the Hawaiian Islands in the late 1700s, and today feral goats occupy a wide variety of habitats on all of the major islands, where they consume native vegetation, trample roots and seedlings, accelerate erosion, and promote the invasion of alien plants. Goats are able to access and forage in extremely rugged terrain, including nearly vertical cliffs of the Nāpali Coast, and they have a high reproductive capacity. Goats can be highly destructive to natural vegetation and contribute to erosion by eating young trees and young shoots of plants before they can become established and by creating trails that can damage native vegetative cover, destabilize substrate, and form gullies that channel water flow and exacerbate erosion. As a result, goats are believed to have completely eliminated some plant species from the Islands. Eradication coupled with major efforts to fence boundaries of Haleakalā and Hawai‘i Volcanoes National Parks have virtually eliminated goat problems, with associated recovery of vegetation. With the development of cattle ranching in the nineteenth and early twentieth centuries, particularly on Hawai‘i, feral cattle became abundant and contributed significantly to the decline of many plant species. Although feral cattle persist today, most cattle grazing is now on managed lands.

8.4 

8.5 

Habitat Impacts from Introduced Animals

Feral pigs in Big Bog, Haleakalā, East Maui

Goats with Diospyros sandwicensis, East Maui

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Ecosystem damage has also been caused by mouflon sheep (Ovis aries orientalis, sometimes considered a separate species as O. musimon), which were deliberately introduced to Kaua‘i, Lāna‘i, and Hawai‘i beginning in the late 1950s by the then Hawai‘i Department of Fish and Wildlife. Food preferences, grazing and browsing behavior, and herding habits are similar to those of feral goats, and native plants are particularly susceptible to damage by mouflon. Efforts to remove these feral animals are now underway on Hawai‘i. Black-tailed deer (mule deer, Odocoileus hemionus) were first introduced to Kaua‘i in 1961 for sport hunting. These deer are currently limited to the western side of Kaua‘i, where they feed on a variety of native and alien plants. Axis deer (Axis axis, from India) have been introduced in the past as game animals on O‘ahu, Moloka‘i, Lāna‘i, and Maui. Maui is currently experiencing a population explosion of axis deer, with substantial damage to both domestic (agriculture) and wild nature. They have recently been illegally brought from Maui to Hawai‘i. In addition to directly impacting native plants through browsing, deer, like pigs, serve as a primary vector for the spread of introduced plants. Deer feed on many alien plant species and distribute seeds of these plants through their feces as they travel. There are 3 species of introduced rats in the Hawaiian Islands. The Polynesian rat (Rattus exulans) probably arrived in the Hawaiian Islands as an inadvertent introduction by early Polynesian colonizers from the central Pacific, while the black rat (Rattus rattus) and the Norway rat (Rattus norvegicus) likely arrived in the Islands as stowaways on nineteenth-century ships. While introduced rats are best known for their impacts on island birds and snails, rat predation on seeds and young plants can seriously affect the regeneration of many native species. Research has indicated that rats may consume as much as 90% of the seeds produced by some species, with plants having large, fleshy fruits being notably susceptible to rat predation. Concern has been raised in particular about rat predation on the fruits of Pritchardia, Cyanea, and Cyrtandra. Although often overlooked, there are a number of alien invertebrate species in the Hawaiian Islands that have had significant impacts on native plant populations, and others that are cause for concern. Predation by nonnative slugs presents a potential threat to all species of Cyanea and perhaps other lobeliads, and a number of species have been observed to be killed in the wild by slug damage. Two insect pests, the black twig borer (Xylosandrus compactus) and the two-spotted leafhopper (Sophonia rufofascia), are relatively recent introductions that attack many native species. Of particular note, however, has been the invasion of yellowjacket wasps (Vespula pensylvanica) to the Hawaiian Islands. These were first reported from Kaua‘i in 1919, but it wasn’t until a particularly aggressive race was introduced in 1977 that scientists began to see yellowjacket wasps expanding their range and dramatically influencing native ecosystems. Although the wasps are diet generalists, they are voracious predators on native insects and spiders. Major concern has been raised at Haleakalā National Park because of the impact that yellowjacket wasps have had on many native insects, including yellow-faced bees (genus Hylaeus), which are the primary pollinator of silverswords. Another insect of concern is the Argentine ant, which became established at Haleakalā in the 1960s and is slowly spreading its

area of invasion. This ant is an overwhelming threat to nonforested high-elevation ecosystems of Haleakalā and Hawai‘i. If it expands its range, it is likely to eliminate many native insect species, including pollinators and numerous flightless local endemics.

Habitat Destruction and Modification by Fire Multiple lines of evidence indicate that the natural fire regime in the Hawaiian Islands before the arrival of man was characterized by infrequent, low-intensity fires. Not only were natural ignitions of fire rare, but moisture levels in the widespread mesic and wet forest habitats were too high to allow for fires to burn easily. In drier forest and shrubland habitats, there was typically an open understory that lacked continuous fuels in which fires could spread. Natural fire regimes, however, began to be altered by a combination of dramatic changes in native land cover as the Polynesians cleared lowland areas for agriculture and the

8.6  Invasion of Andropogon virginicus into mesic forest, Hawai‘i Volcanoes National Park, Hawai‘i

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subsequent arrival and spread of nonnative, fire-adapted grasses associated with the Euro-American colonization in the nineteenth and twentieth centuries. As described in chapter 7, the widespread range expansion of fire-adapted broomsedge (Andropogon virginicus), bushy beard grass (Schizachyrium condensatum), fountain grass (Cenchrus setaceus), and molasses grass (Melinis minutiflora) has dramatically increased the frequency and size of fires in the Hawaiian Islands. These grasses ignite easily and recover quickly after fire, creating a grass-fire cycle and altering native vegetation cover into a new stable state. Most native Hawaiian plants are not adapted to recover well after fire, and there is little or no evidence of Hawaiian species that depend on fire for their existence or regeneration. Only a few native plant species show limited tolerance of fire, including Acacia koa, Dodonaea viscosa, Metrosideros polymorpha, and Osteomeles anthyllidifolia. Higher elevation shrublands have been notably and negatively impacted by fires in recent decades.

Land Use and Land Management Land management in the Hawaiian Islands began with the delineation of ahupua‘a by the ancient Hawaiians. The ahupua‘a was a wedge-shaped area of land running from the uplands to the sea, following the natural boundaries of a watershed, and was managed to maintain a sustainable functioning ecosystem. Each

8.7 

Limahuli Valley, an ahupua‘a on Kaua‘i

ahupua‘a contained the resources that the local village needed, from plants growing in upslope areas to irrigated terraces for farming, to marshy taro fields, and finally to fishponds and the coral reef at the edge of the ocean. Specialized knowledge of resources and technical skills were combined to provide everything needed in the way of food, clothing, and shelter for residents. The ahupua‘a system collapsed for a variety of reasons after the arrival of Euro-American colonists in the nineteenth century, leading to a largely laissez-faire management and exploitation of natural resources. The sandalwood trade provides an early example of this abuse. Trade in this fragrant wood began in the 1790s and expanded rapidly after 1810 into a race to exploit a limited resource. By 1830 the trade had collapsed due to shortage of trees and a drop in prices as new sources of sandalwood were found in India. Modern efforts toward conservation began following the annexation of the Islands by the United States with the creation of the Division of Forestry in 1903, with a major goal of establishing a public-private partnership to protect the upland forests and thereby provide the necessary water resources for lowland agriculture and coastal communities. In 1904, R. S. Hosmer, who had worked with Gifford Pinchot, was appointed Hawaiian superintendent of forestry. The primary focus of Hosmer in the early years of his tenure was the establishment of forest reserves at middle and high elevations, including both government and private lands. His goal was to fence these reserves and remove free-ranging animals such as goats, sheep, cattle, and feral pigs that had already caused serious damage and deforestation. Additionally, he established nurseries that provided trees to landowners for reforestation. Hosmer, following earlier suggestions, distinguished between protection forests on the windward slopes, whose primary function was water yield, and commercial forests on the drier leeward slopes. In 1907, the botanist Joseph Rock was hired to collect plants for introduction to the Hawaiian Islands, an activity he continued until 1920. By the time that Hosmer resigned in 1914 to become chairman of the Department of Forestry at Cornell University, 27 forest reserves had been established covering about 280,000 ha, and efforts had been made to protect these areas. At the same time as Hosmer’s work, the Hawaiian Sugar Planters’ Association (HSPA) hired Harold L. Lyon, who arrived in 1907 to take a position as a plant pathologist. While Lyon was concerned with increasing the genetic breeding of sugarcane and in developing new crops for the Hawaiian Islands, he quickly developed a strong interest in watershed protection. After viewing an area of major dieback in ‘ōhi‘a forest on the north slope of Maui in 1909, Lyon developed the theory that native Hawaiian species were poorly adapted to colonization after disturbance and that further forest decline was inevitable without a major effort at reforestation using non-Hawaiian tree species. These concerns led HSPA to establish its own Department of Agriculture and Forestry. Unsatisfied by efforts of the Division of Forestry, and perhaps in competition with Joseph Rock, Lyon traveled widely, collecting seeds and plants to bring to Hawai‘i. By 1927, he is thought to have introduced as many as 900 tree and shrub species. Because Lyon was concerned with watershed protection, he considered trees with high water use such as many species of eucalyptus as undesirable. Nevertheless, a diverse assemblage

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of eucalypts became some of the most successful plantings. The most commonly planted alien trees in the Hawaiian Islands between 1908 and 1960, in order of numbers planted and combining trees planted for plantations and watershed protections, were the following: Eucalyptus robusta (Myrtaceae), Grevillea robusta (Proteaceae), Melaleuca quinquinervia (Myrtaceae), Casuarina glauca (Casuarinaceae), Cryptomeria japonica (Cupressaceae), Fraxinus uhdei (Oleaceae), Eucalyptus saligna, and Eucalyptus camaldulensis. There were certainly Hawaiian botanists, most notably Otto Degener, who raised objections to the wisdom of introducing so many alien plants to the Hawaiian Islands. Lyon and others, however, remained confident that new tree species would protect watersheds better than native species and that new stable forest communities could be established. Many of the early tree introductions, however, have become invasive alien species of concern and have left a significant legacy of unintended consequences. At the same time that watershed protection was a major program by both the Division of Forestry and HSPA, the Territorial Conservation Commission of Hawai‘i was created in 1908 with the charge to investigate the natural resources of the Islands and recommend their wise development and use. Following this, the Division of Fish and Game was established in 1927 under the Board of Agriculture and Forestry, and that agency has evolved over the intervening years to become today’s Division of Forestry and Wildlife within the Hawai‘i Department of Land and Natural Resources.

Land Management Today Nearly half of the 16,600 km2 total land area of the Hawaiian Islands is managed today by the state or federal governments. The largest landowner is the State of Hawai‘i, with over 4,670 km2 that are managed for a variety of functions, including agricultural use, recreation, transportation, watershed protection, and natural resources protection. An additional 820 km2 are controlled by the Hawai‘i State Department of Hawaiian Home Lands, which manages the lands in trust for present and future use by Native Hawaiians. The federal government, through three agencies (the National Park Service, the U.S. Fish and Wildlife Service, and the Department of Defense), controls more than 2,700 km2 of the Hawaiian Islands. These lands serve a variety of purposes, including not only the conservation of natural and cultural resources and the protection of wildlife habitat but also military support and training and public safety. There are no lands in the Hawaiian Islands managed by either the U.S. Forest Service or the Bureau of Land Management. The remaining Hawaiian land area, forming about 51% of the total, is in private ownership. Combining those lands that receive at least some degree of resource management for conservation, almost a third of the land area of the Hawaiian Islands broadly fits this category of land use. At the state level, there are approximately 105 km2 of land area in 55 state parks, 2,602 km2 in state forest reserves, and 384 km2 in state wildlife sanctuaries. The U.S. National Park Service manages 1,477 km2 of land area, with the largest amount of this in Haleakalā (121 km2) and Hawai‘i Volcanoes National Parks (1,293 km2). Small land areas are designated national wildlife refuges managed by the U.S. Fish and Wildlife Service.

Much of the private land in the Hawaiian Islands is controlled by a small number of large corporations whose ownership dates back to the lands owned or deeded from the Hawaiian royal family in the nineteenth century. Just seven corporations own approximately 20% of the land in the state. These are Kameha­ meha Schools (formerly the Bishop Estate), Parker Ranch, Castle & Cooke, Alexander and Baldwin, the James Campbell Estate, C. Brewer and Company, and Dole Foods. There has been an increasing willingness of some of these owners in recent years to manage significant areas of these lands for watershed protection and conservation purposes in cooperative watershed partnerships with other landowners and state and federal agencies. Rather than conservation easements or land acquisitions, these partnerships are a primary vehicle for conservation on private lands in the Hawaiian Islands. Watershed partnerships are voluntary alliances of public and private landowners that are committed to the common value of protecting forested watersheds for water recharge, conservation, and other ecosystem services through collaborative management. Today there are nine watershed partnerships covering hundreds of thousands of hectares on six islands, and together they involve over forty-five private landowners and twenty-four public agencies in the state. The first watershed partnership began in East Maui in 1991, followed by the West Maui Watershed Partnership (1998), Ko‘olau Watershed Partnership on O‘ahu (1999), East Moloka‘i Watershed Partnership (1999), Lāna‘i Forest and Watershed Partnership (2001), Kaua‘i Watershed Partnership (2003), Haleakalā Watershed Restoration Partnership (2003), Kohala Watershed Partnership (2003), and Three Mountain Alliance (2007). Most management actions by the partnerships to date are habitat based and focus on threats from invasive species and feral ungulates, native habitat restoration, and rare plant protection. A major focus of habitat restoration in the Hawaiian Islands has been the remaining areas of dry forest. The combined impacts of habitat destruction, introduced ungulates, invasive plants, and wildland fires have dramatically reduced the extent and condition of these forests over the past century. Initial restoration programs in the 1980s and 1990s began with the construction of exclosures to exclude ungulates and attempts to remove invasive grass species using both herbicides and hand weeding. Although control of invasive plants requires continued management, the work becomes less intensive as the natural system recovers. Firebreaks, rodent control, and fences to prevent grazing have been found to be critical to protect the recovering forest. While some projects have had mixed results because of the difficulty in providing appropriate microhabitat conditions for seedling germination and establishment, there has been increasing success, giving rise to cautious optimism. These restoration efforts are excellent examples of cooperative collaborations between public and private groups.

Managing Invasive Species While habitat loss through land use changes brought on both by early Polynesian and Euro-American populations have been the major cause of species endangerment and extinction in the past, alien species impacts have now become the predominant

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cause of biodiversity loss in the Hawaiian Islands. These alien species have come from both deliberate and unintentional introductions, with the latter becoming increasingly significant in recent decades given Hawai‘i’s role as a key transportation hub for the Pacific Basin. Invasive alien species not only affect biodiversity and associated ecosystem services, they also have a significant impact on Hawai‘i’s agriculture, business economy, public health, and quality of life. Without question, the impacts from invasive alien species in the Hawaiian Islands far exceed those of any other state in the United States. This situation presents major challenges to federal, state, and local governments. A number of collaborative efforts in recent years have focused on ways to minimize and mitigate damaging impacts from alien species. In 1992, the State of Hawai‘i established the Hawaiian Biological Survey, based at Bishop Museum in Honolulu. The survey provides several useful elements that are updated annually to aid in combating invasive species in Hawai‘i: (1) a comprehensive checklist of species for the state; (2) a list of authoritative

8.8 

Auwahi dry forest restoration plots, East Maui

names to ensure consistent taxonomic usage; (3) the ability to determine whether or not a particular species is known to exist in the Hawaiian Islands and a documentation of its spatial occurrence; and (4) the basis for quick screening of biological control agents by agricultural authorities. In 1996, the U.S. Geological Survey and University of Hawai‘i developed the Hawaiian Ecosystems at Risk project as a grassroots network to share invasive species information for the support and enhancement of the Hawaiian conservation community. Their Web site (www.hear.org) provides a rich link to relevant information. The Hawai‘i Natural Heritage Program of the Nature Conservancy provided a third important effort. Since its establishment in 1985, its focus has moved rare species to the development of a wide range of biodiversity-related spatial data as well as key capabilities in geographical information systems and modeling. Sadly, both of these programs are suffering from a lack of the funding necessary to achieve their goals and may not continue.

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8.9 

Pristine forest landscape, Kaua‘i

Global Change and Hawaiian Ecosystems Island ecosystems are considered among the most vulnerable regions to climate change because of the impact that extreme events can have on them and the sharp environmental gradients that exist. Existing data demonstrates that the climate of the Hawaiian Islands is changing in ways that are consistent with the expected influence of global warming: air temperatures have risen, rainfall and stream flow have decreased, rain intensity has increased, sea level and sea surface temperatures have increased, and the ocean waters are acidifying. The position of the North Pacific subtropical anticyclone (high pressure center) and the altitude of the trade wind inversion are fundamental drivers of local climate and rainfall in the Hawaiian Islands. For this reason, upper-elevation Hawaiian wet forests and similar tropical cloud forests may be among the most sensitive world ecosystems to climate change. Seasonal rainfall variability in the Hawaiian Islands is strongly correlated with fluctuations in the height and strength of the trade wind inversion, and El Niño–Southern Oscillation (ENSO) events are associated with a lower level of this inversion. However, there is not

yet a clear understanding to know whether ENSO activity will be enhanced or damped, or if the frequency of events will change. Lowering the trade wind inversion would reduce rainfall in the Hawaiian Islands in general, but especially at high elevations, and almost certainly depress the upper transition between forest and grassland and heavily impact endangered native forest birds. The driest years in the past century have generally coincided with ENSO events. Warmer air temperatures in the Hawaiian Islands have already had significant adverse affects on native bird species, as warmer conditions expand the range of mosquitoes into higher mountain elevations. These mosquitoes are significant vectors for avian malaria and avian pox, which have led to dramatic declines in native bird populations. Preserving honeycreepers and other native bird species will require active environmental management of those areas they currently inhabit and the elimination or containment of mosquito populations. Warmer air temperatures and drier conditions may also be related to observations of increased mortality and low reproduction rates of silverswords in recent years in Haleakalā National Park. There are now efforts underway to develop and implement a monitoring program to detect changes in species distribution

C O N S E R VA T I O N O F T H E H A W A I I A N F L O R A

in montane wet forest habitats and the upper forest limit. Recent studies on the upper windward slopes of Haleakalā suggest that communities will respond to reductions in moisture balance through changes in species assemblage and structure, especially at the ecotone marking the upper forest line. Climate change forecasts predict increased episodes of heavy storms and hurricane-level winds in the Pacific, with an unknown change in the frequency and strength of ENSO conditions that are often associated with hurricanes. With further warming, hurricane and typhoon peak wind intensities and storm-driven rainfall are likely to increase, which combined with sea level rise would cause higher storm surge levels. These changes would be expected to have major impacts for natural habitats, species, and resources. The scientific evidence for future sea level rise is strong and unequivocal. Low-lying coastal areas will face increased risk from sea water inundation due to sea level rise and storm surge, with major consequences for coastal communities, infrastructure, natural habitats, and resources. Research currently indicates that global mean sea level rise may reach approximately 30 cm by 2050, and best current models predict an increase of 60-100 cm over the next century. There are, however, significant unknowns in predicting future sea level.. While coastal areas and beaches of Hawai‘i are relatively protected today from storm surges by their well-developed coral reefs, these reefs are expected to be sensitive to global changes in the form of ocean acidification and temperature increase that could significantly impact reef structure. Without the protective services of these coral reefs, sandy beaches and coastline areas will likely undergo increased rates of erosion. Sea level rise has special meaning for the low-lying atolls and sandy shoals of the Papahānaumokuākea Marine National Monument, where coastal erosion and heightened damage from storm surge, high wave events, extreme tides, and tsunami inundation are likely in the future. The biota of this area may be greatly impacted from rises in sea level because of the nesting and nursing habitats of seabirds and concentrations of endangered and threatened species. Hurricanes have been relatively rare events in the Hawaiian Islands, but models suggest that their frequency and intensity may likely increase in the future. Five major hurricanes have struck the Hawaiian Islands and caused serious damage since 1950. Hurricane Nina (1957) produced record winds in Honolulu, Hurricane Dot (1959) caused damage to Kaua‘i, Hurricane

Estelle (1986) produced floods on O‘ahu and very high surf on Hawai‘i and Maui, and Hurricane Iwa (1982) caused hundreds of millions of dollars of damage on Kaua‘i. Kaua‘i also received the brunt of Hurricane ‘Iniki (1992), which struck the island with sustained winds of 200 kph and caused over $2.3 billion in property damage. The Hawaiian terrain makes the islands particularly sensitive to damage from hurricanes, which focus torrential rains on mountain slopes, resulting in destructive flash floods and landslides. The State of Hawai‘i has recognized that it is critical to act now in order to prepare for the impacts of climate change to better withstand the negative impacts and take advantage of positive opportunities. The approach being developed has involved an open and collaborative planning process that is based on establishing a common vision for the future of the Hawaiian Islands that remains flexible to changing environment, economy, and society. Working proactively in this way will help to ensure that Hawai‘i’s future is one in which the Hawaiian population and its rich biological diversity will remain healthy and resilient in the face of global change.

Conclusions Much of the conservation effort in the Hawaiian Islands to date has been focused on the protection of rare and endangered species. While such work with rare species is crucial, much more needs to be done to protect functioning ecosystems and their associated species and ecosystem services. The highest priorities in this respect are the control of invasive alien species and, more specifically, the elimination of feral pig damage. Without these steps, it will be difficult if not impossible to provide for the effective restoration of damaged habitats and preserve the remaining native Hawaiian biodiversity. Issues of global change will complicate these efforts. There is considerable optimism that much can be accomplished if the political will exists at both the local and state level to implement the needed management. More research is needed, but at the same time the critical factor for conservation is the need for a public appreciation for the value of the biodiversity heritage. There is concern, however, that public apathy may represent the greatest threat for the future of conservation in the Hawaiian Islands. Public outreach and education will do much to determine the future.

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APPENDIX

Native Vascular Plants of the Hawaiian Islands

Warren L. Wagner (Smithsonian Institution, Department of Botany), Nancy R. Khan (Smithsonian Institution, Department of Botany), David H. Lorence (National Tropical Botanical Garden), and Derral R. Herbst (Bishop Museum)

Ml Mo N Ne Ni NWI O PH Wa WM

The following list is comprised of the 1,352 vascular plant taxa considered to be native to seventeen main islands in the Hawaiian Archipelago. Plants are grouped in three main categories: ferns and fern allies, monocots, and dicots. Within each group, families are arranged alphabetically as are the individual genera and species. Specific information about the nativity, distribution, and conservation status of each taxon is provided based on the following abbreviations:

Molokini Island Moloka‘i Island Nīhoa Island Necker Island Ni‘ihau Island Northwestern Hawaiian Islands O‘ahu Island Pearl and Hermes Atoll Wai‘anae Mountains, O‘ahu West Maui

Conservation Assessment (Wagner et al., 1999) EX extinct EXH extirpated in the Hawaiian Islands E endangered V vulnerable R rare

Nativity: end endemic ind indigenous Note: Under the island distribution of some species, where it is not clear whether it arrived naturally to the Hawaiian Islands and hence is indigenous, or whether it was brought more recently with the assistance of humans, we have added “pol?” to indicate that it is questionable and may have been human introduced.

US Status (U.S. Fish and Wildlife Service) E endangered T threatened C candidate

Distribution: EM East Maui FF French Frigate Shoals (incl. Tern and other islands) GP Gardner Pinnacles H Hawai‘i Island HI The eight main Hawaiian Islands (Ni‘ihau, Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, Maui, Kaho‘olawe, and Hawai‘i) K Kaua‘i Island Ka Kaho‘olawe Island Kl Ka‘ūla Island Ko Ko‘olau Mountains, O‘ahu Ku Kure Atoll (incl. Sand and Green Islands) L Lāna‘i Island La Laysan Island Le Lehua Island Li Lisianski Island M Maui Island Mi Midway Atoll (incl. Sand and Eastern Islands)

This list was generated from data assembled for the Flora of the Hawaiian Islands Web site (http://botany.si.edu/pacificisland​ biodiversity/hawaiianflora/index.htm), which is a collaboration between the Smithsonian Institution and the National Tropical Botanical Garden (NTBG). Database management and technical editing was handled by Nancy Khan, with scientific oversight and content review provided by Warren L. Wagner, David H. Lorence, and Derral R. Herbst. Valuable expertise and comments were also provided by Tim Flynn and Ken Wood, botanists at NTBG. Wagner, W. L., M. Bruegmann, D. R. Herbst, and J. Q. Lau. 1999. Hawaiian vascular plants at risk: 1999. Bishop Museum Occasional Papers 60: 1–58. Wagner, W. L., D. R. Herbst, and S. H. Sohmer. 1999. Manual of the Flowering Plants of Hawai‘i. Revised edition with supplement by W. L. Wagner and D. R. Herbst, pp. 1855–1918. University of Hawai‘i Press, Honolulu, 1919 pp. in 2 volumes. (Bishop Museum Special Publication 97).

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FERNS AND FERN ALLIES

Aspleniaceae Asplenium acuminatum Hook. & Arn. end; HI exc. Ni, Ka Asplenium adiantum-nigrum L. ind; HI exc. Ni, Ka Asplenium aethiopicum (Burm. f.) Bech. ind; HI exc. Ni, Ka Asplenium caudatum G. Forest. ind; HI exc. Ni, Ka Asplenium contiguum Kaulf. var. contiguum end; HI exc. Ni, Ka var. hirtulum C. Chr. end; K/ M Asplenium dielerectum Viane end; O (southern Ko‘olau)/ Mo/ L (ex)/ M/ H (Manukā and Honomalino); Conservation Assessment = E; US Status = E Asplenium dielfalcatum Viane end; O (Wa and Ko (ex)); Conservation Assessment = E; US Status = E Asplenium diellaciniatum Viane end; K (Kawai‘iki)/; Conservation Assessment = E; US Status = E Asplenium dielmannii Viane end; K (Halemanu); Conservation Assessment = E; US Status = E Asplenium dielpallidum N. Snow end; K (now only Koai‘e, Ku‘ia, Mahanaloa, Mākaha); Conservation Assessment = E; US Status = E Asplenium glabratum W. J. Rob. end; HI exc. Ni, Ka Asplenium haleakalense W. H. Wagner end; EM (Waikamoi, Kīpahulu, Wai‘ānapanapa)/ H (Maka‘ala Natural Area Reserve) Asplenium hobdyi W. H. Wagner end; K/ Mo/ M/ H Asplenium insiticium Brack. ind; HI exc. Ni, Ka Asplenium kaulfussii Schltdl. end; K/ O/ Mo/ L/ M/ H Asplenium leucostegioides Baker end; EM (extinct); Conservation Assessment = EX Asplenium lobulatum Mett. ind; HI exc. Ni, Ka Asplenium macraei Hook. & Grev. end; HI exc. Ni, Ka Asplenium monanthes L. ind; M/ H Asplenium nidus L. ind; HI exc. Ni, Ka Asplenium normale D. Don ind; HI exc. Ni, Ka Asplenium peruvianum Desv. var. insulare (C. V. Morton) D. D. Palmer end; M/ H; Conservation Assessment = E; US Status = E

Asplenium polyodon G. Forst. ind; HI exc. Ni, Ka Asplenium schizophyllum C. Chr. end; K (Kōke‘e area)/ H; Conservation Assessment = R Asplenium sphenotomum Hillebr. end; HI exc. Ni, Ka Asplenium trichomanes L. subsp. densum (Brack.) W. H. Wagner end; EM/ H Asplenium unisorum (W. H. Wagner) Viane end; O (southern Wa); Conservation Assessment = E; US Status = E Hymenasplenium unilaterale (Lam.) Hayata ind; HI exc. Ni, Ka Blechnaceae Doodia kunthiana Gaudich. end; HI exc. Ni, Ka Doodia lyonii O. Deg. end; K (Koai‘e)/ O/ M/ H (ex); Conservation Assessment = V Sadleria cyatheoides Kaulf. end; HI exc. Ni, Ka Sadleria pallida Hook. & Arn. end; HI exc. Ni, Ka Sadleria souleyetiana (Gaudich.) T. Moore end; HI exc. Ni, Ka Sadleria squarrosa (Gaudich.) T. Moore end; HI exc. Ni, Ka Sadleria unisora (Baker) W. J. Rob. end; K Sadleria wagneriana D. D. Palmer & Flynn end; K (Kauaikananā and Mōhihi Streams, and Wainiha pali) Cibotiaceae Cibotium chamissoi Kaulf. end; O/ Mo/ L/ M/ H Cibotium glaucum (Sm.) Hook. & Arn. end; HI exc. Ni, Ka Cibotium menziesii Hook. end; HI exc. Ni, Ka Cibotium nealiae O. Deg. end; K Dennstaedtiaceae Hypolepis hawaiiensis Brownsey var. hawaiiensis end; HI exc. Ni, Ka var. mauiensis (Hillebr.) D. D. Palmer end; M Microlepia speluncae (L.) T. Moore ind; K/ O/ Mo/ WM/ H Microlepia strigosa (Thunb.) C. Presl var. mauiensis (W. H. Wagner) D. D. Palmer end; M/ H; Conservation Assessment = R; US Status = C var. strigosa ind; HI exc. Ni, Ka Pteridium aquilinum (L.) Kuhn subsp. decompositum (Gaudich) Lamoureux ex J. A. Thomson end; HI exc. Ni, Ka

APPENDIX

Dryopteridaceae Arachniodes insularis W. H. Wagner end; K/ O/ Mo/ M/ H (Kohala Mts) Ctenitis latifrons (Brack.) Copel. end; HI exc. Ni, Ka Ctenitis squamigera (Hook. & Arn.) Copel. end; K (Kōke‘e)/ O/ Mo/ L/ M; Conservation Assessment = E; US Status = E Cyrtomium caryotideum (Wall.) C. Presl ind; HI exc. Ni, Ka Dryopteris crinalis (Hook. & Arn.) C. Chr. var. crinalis end; K/ O/ Mo/ L/ M/ H var. podosorus (W. H. Wagner & Flynn) D. D. Palmer end; K (Kōke‘e; Kawaikōī); Conservation Assessment = R; US Status = E Dryopteris fuscoatra (Hillebr.) W. J. Rob. var. fuscoatra end; HI exc. Ni, Ka var. lamoureuxii Fraser-Jenk. end; M (Makawao and Ko‘olau Forest Reserves, Kaua‘ula Valley); Conservation Assessment = R Dryopteris glabra (Brack.) Kuntze var. alboviridis (W. H. Wagner) D. D. Palmer end; K (Kōke‘e, Wai‘ale‘ale, and Alakai); Conservation Assessment = R var. flynnii D. D. Palmer end; K (Pihea); Conservation Assessment = R var. glabra end; HI exc. Ni, Ka var. hobdyana (W. H. Wagner) D. D. Palmer end; EM (north slope) var. nuda (Underw.) Fraser-Jenk. end; K/ O/ Mo/ M var. pusilla (Hillebr.) Fraser-Jenk. end; K (Kōke‘e); perhaps on other islands var. soripes (Hillebr.) Heart ex Fraser-Jenk. end; K/ O/ Mo/ M/ H Dryopteris hawaiiensis (Hillebr.) W. J. Rob. end; K/ O/ Mo/ M/ H Dryopteris mauiensis C. Chr. end; HI exc. Ni, Ka Dryopteris rubiginosa (Brack.) Kuntze end; HI exc. Ni, Ka Dryopteris sandwicensis (Hook. & Arn.) C. Chr. end; HI exc. Ni, Ka Dryopteris subbipinnata W. H. Wagner & Hobdy end; M/ H Dryopteris tetrapinnata W. H. Wagner & Hobdy end; EM; Conservation Assessment = R Dryopteris unidentata (Hook. & Arn.) C. Chr. var. paleacea (Hillebr.) Heart ex Fraser-Jenk. end; K/ O/ Mo/ M/ H var. unidentata end; HI exc. Ni, Ka Dryopteris wallichiana (Spreng.) Hyl. ind; K/ O/ Mo/ M/ H

Elaphoglossum aemulum (Kaulf.) Brack. end; K/ O/ Mo/ L/ M/ H (ex) Elaphoglossum alatum Gaudich. end; O Elaphoglossum crassicaule Copel. end; K Elaphoglossum crassifolium (Gaudich.) W. R. Anderson & Crosby end; HI exc. Ni, Ka Elaphoglossum fauriei Copel. end; O (Ko)/ Mo Elaphoglossum paleaceum (Hook. & Grev.) Sledge ind; K/ O/ Mo/ L (Kunoa Gulch)/ M/ H Elaphoglossum parvisquameum Skottsb. end; Mo/ L/ M/ H Elaphoglossum pellucidum Gaudich. end; K/ O/ Mo/ M/ H Elaphoglossum wawrae (Luerss.) C. Chr. end; K/ O/ Mo/ M/ H Polystichum bonseyi W. H. Wagner & Hobdy end; EM/ H (single collection on Hualālai) Polystichum haleakalense Brack. end; EM/ H (Mauna Kea) Polystichum hillebrandii Carruth. end; M/ H Gleicheniaceae Dicranopteris linearis (Burm. f.) Underw. ind; HI exc. Ni, Ka Diplopterygium pinnatum (Kunze) Nakai ind; HI exc. Ni, Ka Sticherus owhyhensis (Hook.) Ching end; HI exc. Ni, Ka Hymenophyllaceae Callistopteris baldwinii (D. C. Eaton) Copel. end; HI exc. Ni, Ka Crepidomanes draytonianum (Brack.) Ebihara & K. Iwats. end; HI exc. Ni, Ka Crepidomanes minutum (Blume) K. Iwats. ind; HI Hymenophyllum lanceolatum Hook. & Arn. end; HI exc. Ni, Ka Hymenophyllum obtusum Hook. & Arn. end; O/ Mo/ L/ M/ H Hymenophyllum recurvum Gaudich. end; HI exc. Ni, Ka Vandenboschia cyrtotheca (Hillebr.) Copel. end; HI exc. Ni, Ka Vandenboschia davallioides (Gaudich.) Copel. end; HI exc. Ni, Ka Vandenboschia tubiflora F. S. Wagner end; K Isoëtaceae Isoëtes hawaiiensis W. C. Taylor & W. H. Wagner end; WM (‘Eke)/ H (Mauna Kea); Conservation Assessment = R

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Lindsaeaceae Lindsaea repens (Bory) Thwaites var. macraeana (Hook. & Arn.) C. Chr. end; HI exc. Ni, Ka Sphenomeris chinensis (L.) Maxon ind; HI exc. Ni, Ka Lomariopsidaceae Nephrolepis cordifolia (L.) C. Presl ind; HI exc. Ni, Ka Nephrolepis exaltata (L.) Schott subsp. hawaiiensis W. H. Wagner end; HI exc. Ka Lycopodiaceae Huperzia erosa Beitel & W. H. Wagner end; HI exc. Ni, Ka Huperzia erubescens (Brack.) Holub ind; K/ O/ Mo/ M/ H Huperzia filiformis (Sw.) Holub ind; K/ O/ Mo/ L/ M/ H Huperzia haleakalae (Brack.) Holub ind; EM (extinct); Conservation Assessment = EXH Huperzia mannii (Hillebr.) Kartesz & Gandhi end; K (ex)/ M/ H; Conservation Assessment = E; US Status = E Huperzia nutans (Brack.) Rothm. end; K (Wahiawa Mts, ex)/ O (Ko); Conservation Assessment = E; US Status = E Huperzia phyllantha (Hook. & Arn.) Holub ind; HI exc. Ni, Ka Huperzia serrata (Thunb. ex Murray) Trevis. ind; K/ O/ Mo/ L/ H Huperzia stemmermanniae (A. C. Medeiros, W. H. Wagner & Hobdy) Kartesz end; EM/ H; Conservation Assessment = E; US Status = C Huperzia subintegra (Hillebr.) Beitel & W. H. Wagner end; K/ O/ Mo/ M Lycopodiella cernua (L.) Pic. Serm. ind; HI exc. Ni, Ka Lycopodium venustulum Gaudich. var. venustulum ind; HI exc. Ni, Ka var. verticale W. H. Wagner end; H (Mauna Kea) Marattiaceae Marattia douglasii (C. Presl) Baker end; HI exc. Ni, Ka Marsileaceae Marsilea villosa Kaulf. end; Ni/ O/ Mo; Conservation Assessment = E; US Status = E Ophioglossaceae Botrychium subbifoliatum Brack. end; HI exc. Ni, Ka (extinct); Conservation Assessment = EX

Ophioglossum nudicaule L. f. ind; K/ O/ H (Kīlauea) Ophioglossum pendulum L. ind; K/ O (Ko)/ Mo (Wailau Valley)/ L/ M/ H Ophioglossum petiolatum Hook. ind; K/ O/ Mo (Kaunakakai)/ L/ M/ H Ophioglossum polyphyllum A. Braun ind; K/ O/ Mo (Mo‘omomi)/ L (ex)/ M/ Ka/ H Polypodiaceae Adenophorus abietinus (D. C. Eaton) K. A. Wilson end; K/ O (Ko) / L/ M Adenophorus epigaeus (L. E. Bishop) W. H. Wagner end; K Adenophorus haalilioanus (Brack.) K. A. Wilson end; K (N part and Wahiawa Bog area)/ O (Ko) Adenophorus hymenophylloides (Kaulf.) Hook. & Grev. end; HI exc. Ni, Ka Adenophorus oahuensis (Copel.) L. E. Bishop end; O (Ko) Adenophorus periens L. E. Bishop end; K/ O (ex)/ Mo/ L (ex)/ M (ex)/ H (Glenwood only extant); Conservation Assessment = E; US Status = E Adenophorus pinnatifidus Gaudich. var. pinnatifidus end; HI exc. Ni, Ka var. rockii (Copel.) D. D. Palmer end; K/ O (Ko)/ Mo/ M Adenophorus tamariscinus (Kaulf.) Hook. & Grev. var. montanus (Hillebr.) L. E. Bishop end; Mo/ M/ H (Kohala Mts) var. tamariscinus end; HI exc. Ni, Ka Adenophorus tenellus (Kaulf.) Ranker end; HI exc. Ni, Ka Adenophorus tripinnatifidus Gaudich. end; HI exc. Ni, Ka Grammitis baldwinii (Baker) Copel. end; K Grammitis forbesiana W. H. Wagner end; K/ Mo/ M Grammitis hookeri (Brack.) Copel. ind; HI exc. Ni, Ka Lepisorus thunbergianus (Kaulf.) Ching ind; HI exc. Ni, Ka Microsorum spectrum (Kaulf.) Copel. var. pentadactylum (Hillebr.) D. D. Palmer end; K/ EM (Nāhiku) var. spectrum end; HI exc. Ni, Ka Polypodium pellucidum Kaulf. var. acuminatum D. D. Palmer end; K (Waimea and Hanalei)/ EM (Kaupō Gap) var. pellucidum end; HI exc. Ni, Ka var. vulcanicum Skottsb. end; Mo/ M/ H Stenogrammitis saffordii (Maxon) Labiak end; HI exc. Ni, Ka

APPENDIX

Psilotaceae Psilotum complanatum Sw. ind; HI exc. Ni, Ka Psilotum nudum (L.) P. Beauv. ind; Mi/ HI Pteridaceae Adiantum capillus-veneris L. ind; Ni/ K/ O/ Mo/ L/ M/ H Coniogramme pilosa (Brack.) Hieron. end; HI exc. Ni, Ka Doryopteris angelica K. R. Wood & W. H. Wagner end; K (Kōke‘e); US Status = E Doryopteris decipiens (Hook.) J. Sm. end; HI Doryopteris decora Brack. end; HI exc. Ni Doryopteris takeuchii (W. H. Wagner) W. H. Wagner end; O (Diamond Head); Conservation Assessment = R; US Status = E Haplopteris elongata (Sw.) E. H. Crane ind; HI exc. Ni, Ka Pellaea ternifolia (Cav.) Link ind; HI exc. Ni, Ka Pteris cretica L. ind; HI exc. Ni, Ka Pteris excelsa Gaudich. ind; HI exc. Ni, Ka Pteris hillebrandii Copel. end; HI exc. Ni, Ka Pteris irregularis Kaulf. end; HI exc. Ni, Ka Pteris lidgatei (Baker) H. Christ end; O (Ko)/ Mo / WM; Conservation Assessment = E; US Status = E Schizaeaceae Schizaea robusta Baker end; HI exc. Ni, Ka Selaginellaceae Selaginella arbuscula (Kaulf.) Spring end; HI exc. Ni, Ka Selaginella deflexa Brack. end; K/ O/ Mo/ M/ H Tectariaceae Tectaria gaudichaudii (Mett.) Maxon end; HI exc. Ni, Ka Thelypteridaceae Cyclosorus boydiae (D. C. Eaton) W. H. Wagner end; O (Ko)/ EM (Kīpahulu and Waiho‘i valleys); Conservation Assessment = R; US Status = C Cyclosorus cyatheoides (Kaulf.) Farw. end; HI exc. Ni, Ka Cyclosorus hudsonianus (Brack.) Ching end; HI exc. Ni, Ka

Cyclosorus interruptus (Willd.) H. Itō ind; HI exc. Ni, Ka Cyclosorus pendens (D. D. Palmer) N. Snow end; K/ O/ Mo/ M/ H Cyclosorus sandwicensis (Brack.) Copel. end; HI exc. Ni, Ka Cyclosorus wailele (Flynn) W. H. Wagner end; K (waterfalls in Hanalei and Waimea districts); Conservation Assessment = R Pseudophegopteris keraudreniana (Gaudich.) Holttum end; HI exc. Ni, Ka Thelypteris globulifera (Brack.) C. F. Reed end; HI exc. Ni, Ka Woodsiaceae Athyrium microphyllum (Sm.) Alston end; HI exc. Ni, Ka Cystopteris douglasii Hook. end; EM/ H; Conservation Assessment = R Cystopteris sandwicensis Brack. end; K/ O/ L/ WM Deparia cataracticola M. Kato end; K Deparia fenzliana (Luerss.) M. Kato end; HI exc. Ni, Ka Deparia kaalaana (Copel.) M. Kato end; K (Ka‘ala District)/ EM (Kīpahulu)/ H (Hilo and Laupāhoehoe) (extinct); Conservation Assessment = EX Deparia marginalis (Hillebr.) M. Kato end; K/ Mo/ L/ M/ H Deparia prolifera (Kaulf.) Hook. & Grev. end; HI exc. Ni, Ka Diplazium arnottii Brack. end; HI exc. Ni, Ka Diplazium molokaiense W. J. Rob. end; K(ex)/ O(ex)/ Mo(ex)/ L(ex)/ M (now only extant at Honomanū, Polipoli, and Pūehuehu Nui); Conservation Assessment = E; US Status = E Diplazium sandwichianum (C. Presl) Diels end; HI exc. Ni, Ka DICOTS

Aizoaceae Sesuvium portulacastrum (L.) L. ind; Mi/ PH/ Li/ La/ Ne/ HI Amaranthaceae Achyranthes atollensis H. St. John end; Ku/ Mi/ PH/ La (extinct); Conservation Assessment = EX Achyranthes mutica A. Gray end; K (ex)/ H (South Kohala District); Conservation Assessment = E; US Status = E Achyranthes splendens Mart. ex Moq. var. rotundata Hillebr. end; O/ Mo (ex)/ L (ex); Conservation Assessment = E; US Status = E

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var. splendens end; L/ M; Conservation Assessment = R Amaranthus brownii Christoph. & Caum end; N; Conservation Assessment = E; US Status = E Charpentiera densiflora Sohmer end; K (Ho‘olulu; Hanakāpī‘ai; and Hanakoa valleys; Nāpali Coast); Conservation Assessment = V; US Status = E Charpentiera elliptica (Hillebr.) A. Heller end; K Charpentiera obovata Gaudich. end; K/ O/ Mo/ L/ M/ H Charpentiera ovata Gaudich. var. niuensis Sohmer end; O (Ko: Niu and Wailupe valleys) var. ovata end; O (south Ko)/ Mo/ M/ H Charpentiera tomentosa Sohmer var. maakuaensis Sohmer end; O (Ko: windward gulches) var. tomentosa end; O (Wa)/ Mo/ L/ M/ H Chenopodium oahuense (Meyen) Aellen end; Li/ La/ FF/ Ne/ N/ HI Nototrichium divaricatum Lorence end; K (Nāpali); Conservation Assessment = R Nototrichium humile Hillebr. end; O (Wa)/ EM (Luala‘ilua Hills); Conservation Assessment = E; US Status = E Nototrichium sandwicense (A. Gray) Hillebr. end; HI Anacardiaceae Rhus sandwicensis A. Gray end; K (Kamali‘i Ridge; Makaweli)/ O (Ko: Nu‘uanu and Moanalua valleys; Ho‘omaluhia)/ Mo (Pelekunu Valley)/ M/ H Apiaceae Peucedanum sandwicense Hillebr. end; K/ O (Wa)/ Mo/ WM/ EM (Keōpuka Islet); Conservation Assessment = V; US Status = T Sanicula kauaiensis H. St. John end; K (Wai‘ale‘ale; Kalalau Valley) (extinct?); Conservation Assessment = EX? Sanicula mariversa Nagata & S. M. Gon end; O (Wa: ‘Ōhikilolo); Conservation Assessment = E; US Status = E Sanicula purpurea H. St. John & Hosaka end; O (Ko)/ WM; Conservation Assessment = E; US Status = E Sanicula sandwicensis A. Gray end; EM (slopes of Haleakalā)/WM (‘Īao Valley)/ H (Mauna Kea; Mauna Loa; Hualālai); Conservation Assessment = R Spermolepis hawaiiensis H. Wolff end; K/ O/ Mo / L/ M/ H; Conservation Assessment = E; US Status = E

Apocynaceae Alyxia stellata (J. R. Forst. & G. Forst.) Roem. & Schult. ind; HI; no longer occurring on Ni and Ka Ochrosia compta K. Schum. end; O/ Mo (Pelekunu Trail; Wailau Valley; Kahanui)/ WM (Kahakuloa); Conservation Assessment = R Ochrosia haleakalae H. St. John end; EM/ H (Pololū Valley; Kalōpā Gulch); Conservation Assessment = R; US Status = C Ochrosia kauaiensis H. St. John end; K (Nāpali Coast; Hi‘i Mts); Conservation Assessment = R Ochrosia kilaueaensis H. St. John end; H (Pu‘uwa‘awa‘a; Kīpukapuaulu) (extinct); Conservation Assessment = EX; US Status = E Pteralyxia kauaiensis Caum end; K (Ku‘ia, Kalalau, Mahanaloa valleys; Hi‘i Mts; Hā‘upu; Pāpa‘a; Wai‘oli-Waipā Ridge; Limahuli); Conservation Assessment = E; US Status = E Pteralyxia macrocarpa (Hillebr.) K. Schum. end; O; Conservation Assessment = V; US Status = E Rauvolfia sandwicensis A. DC. end; Ni/ K/ O/ Mo/ L/ M/ H Aquifoliaceae Ilex anomala Hook. & Arn. ind; K/ O/ Mo/ L/ M/ H Araliaceae Cheirodendron dominii Krajina end; K (Mt Wai‘ale‘ale; Nāmolokama Plateau); Conservation Assessment = R Cheirodendron fauriei Hochr. end; K (Kōke‘e; Mt Kāhili; Hā‘upu Ridge; Powerline Trail) Cheirodendron forbesii (Sherff) Lowry end; K (Mt Kāhili; Kamo‘oloa Stream; Makaleha Mts; Powerline Trail) Cheirodendron platyphyllum (Hook. & Arn.) Seem. subsp. kauaiense (Krajina) Lowry end; K subsp. platyphyllum end; O Cheirodendron trigynum (Gaudich.) A. Heller subsp. helleri (Sherff) Lowry end; K subsp. trigynum end; Ni/ O/ Mo/ L/ M/ H Polyscias bisattenuata (Sherff) Lowry & G. M. Plunkett end; K (from Pu‘u Kolo to eastern slope of Mt Kāhili); Conservation Assessment = E: US Status = E Polyscias flynnii (Lowry & K. R. Wood) Lowry & G. M. Plunkett end; K (Kalalau); Conservation Assessment = E; US Status =E Polyscias gymnocarpa (Hillebr.) Lowry & G. M. Plunkett end; O (Ko); Conservation Assessment = E; US Status = E Polyscias hawaiensis (A. Gray) Lowry & G. M. Plunkett end; Mo/ L/ M/ H Polyscias kavaiensis (H. Mann) Lowry & G. M. Plunkett end; K/ O (north Wa)/ L/ M/ H

APPENDIX

Polyscias lydgatei (Hillebr.) Lowry & G. M. Plunkett end; O (Ko‘olau Range); Conservation Assessment = E; US Status = E Polyscias oahuensis (A. Gray) Lowry & G. M. Plunkett end; K/ O/ Mo/ L/ M/ H Polyscias racemosa (C. N. Forbes) Lowry & G. M. Plunkett end; K (Nounou Mts; Nāpali Coast; Ha‘ūpu Ridge; Koai‘e Canyon); Conservation Assessment = E; US Status = E Polyscias sandwicensis (A. Gray) Lowry & G. M. Plunkett end; Ni/ O/ Mo/ L/ M/ H; Conservation Assessment = R Polyscias waialealae (Rock) Lowry & G. M. Plunkett end; K Polyscias waimeae (Wawra) Lowry & G. M. Plunkett end; K Asteraceae Adenostemma viscosum J. R. Forst. & G. Forst. ind; K/ O/ Mo/ L/ M/ H Argyroxiphium caliginis C. N. Forbes end; WM (‘Eke; Pu‘ukukui); Conservation Assessment = R Argyroxiphium grayanum (Hillebr.) O. Deg. end; WM summit; EM (above Hāna Forest Reserve) Argyroxiphium kauense (Rock & M. Neal) O. Deg. & I. Deg. end; H (Mauna Loa; Hualālai single collection); Conservation Assessment = E; US Status = E Argyroxiphium sandwicense DC. subsp. macrocephalum (A. Gray) Meyrat end; EM (Haleakalā); Conservation Assessment = V; US Status = T subsp. sandwicense end; H (Mauna Kea); Conservation Assessment = E; US Status = E Argyroxiphium virescens Hillebr. end; EM (Ukulele; Pu‘unianiau; Ko‘olau Gap; Kuiki) (extinct); Conservation Assessment = EX Artemisia australis Less. end; HI Artemisia kauaiensis (Skottsb.) Skottsb. end; K Artemisia mauiensis (A. Gray) Skottsb. end; EM (Haleakalā Crater and south slope) Bidens amplectens Sherff end; O (windward Wa: between Kawaihāpai and Ka‘ena Pt); Conservation Assessment = R; US Status = E Bidens asymmetrica (H. Lév.) Sherff end; O (southeast leeward Ko) Bidens campylotheca Sch. Bip. subsp. campylotheca end; K/ O/ L / H; Conservation Assessment = V subsp. pentamera (Sherff) Ganders & Nagata end; M (WM: only Pāpalaua Gulch); Conservation Assessment = R; US Status = PE subsp. waihoiensis H. St. John end; EM; Conservation Assessment = R; US Status = PE Bidens cervicata Sherff end; Ni/ northwest K/ O (northwest Wa) Bidens conjuncta Sherff end; WM; Conservation Assessment = R; US Status = PE

Bidens cosmoides (A. Gray) Sherff end; K; Conservation Assessment = V Bidens forbesii Sherff subsp. forbesii end; K subsp. kahiliensis Ganders & Nagata end; K (Mt Kāhili; Kapalaoa) Bidens hawaiensis A. Gray end; H (Kohala; Puna District; Kīlauea) Bidens hillebrandiana (Drake) O. Deg. subsp. hillebrandiana end; H (Kohala Coast); Conservation Assessment = V; US Status = PE subsp. polycephala Nagata & Ganders end; Mo (windward coast)/ EM (windward coast) Bidens macrocarpa (A. Gray) Sherff end; O (Ko) Bidens mauiensis (A. Gray) Sherff end; L/ M/ Ka Bidens menziesii (A. Gray) Sherff subsp. filiformis (Sherff) Ganders & Nagata end; H (leeward sides of and in saddle between Mauna Loa and Mauna Kea) subsp. menziesii end; leeward Mo/ leeward WM Bidens micrantha Gaudich. subsp. ctenophylla (Sherff) Ganders & Nagata end; H (leeward Hualālai); Conservation Assessment = V; US Status = PE subsp. kalealaha Ganders & Nagata end; L/ WM (single locality)/ EM; Conservation Assessment = E; US Status = E subsp. micrantha end; WM/ EM (single collection) Bidens molokaiensis (Hillebr.) Sherff end; O (Diamond Head, ex)/ windward Mo; Conservation Assessment = V Bidens populifolia Sherff end; O (windward Ko: between Kaipapa‘u and Ka‘a‘awa); Conservation Assessment = R Bidens sandvicensis Less. subsp. confusa Nagata & Ganders end; K (Waimea Canyon); Conservation Assessment = R subsp. sandvicensis end; K/ O Bidens torta Sherff end; O (Wa; northwest Ko) Bidens valida Sherff end; K (Mt Kāhili; Hā‘upu Ridge) Bidens wiebkei Sherff end; northeast Mo; Conservation Assessment = E; US Status = E Dubautia arborea (A. Gray) D. D. Keck end; H; Conservation Assessment = R Dubautia carrii B. G. Baldwin & Friar end; Mo (Pu‘u Ka‘eha); Conservation Assessment = V Dubautia ciliolata (DC.) D. D. Keck subsp. ciliolata end; H (Hualālai; Kīlauea; Mauna Loa)

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subsp. glutinosa G. D. Carr end; H (Mauna Kea) Dubautia hanaulaensis B. G. Baldwin end; WM (Ukumehame rim); Conservation Assessment = E Dubautia herbstobatae G. D. Carr end; O (Wa: ‘Ōhikilolo; Kamaile‘unu Ridge); Conservation Assessment = E; US Status = E Dubautia imbricata H. St. John & G. D. Carr subsp. acronaea G. D. Carr end; K (Wai‘ale‘ale); Conservation Assessment = R subsp. imbricata end; K (Wahiawa Mts); Conservation Assessment = R; US Status = E Dubautia kalalauensis B. G. Baldwin & G. D. Carr end; K (Kalalau rim); Conservation Assessment = E; US Status = E Dubautia kenwoodii G. D. Carr end; K (Kalalau rim) (extinct?); Conservation Assessment = EX?; US Status = E Dubautia knudsenii Hillebr. subsp. filiformis G. D. Carr end; K; Conservation Assessment = R subsp. knudsenii end; K subsp. nagatae (H. St. John) G. D. Carr end; K Dubautia laevigata A. Gray end; K Dubautia latifolia (A. Gray) D. D. Keck end; K (Kōke‘e); Conservation Assessment = E; US Status = E Dubautia laxa Hook. & Arn. subsp. bryanii (Sherff) G. D. Carr end; O (Ko) subsp. hirsuta (Hillebr.) G. D. Carr end; K/ O (primarily Wa)/ L subsp. laxa end; K (Hā‘upu summit)/ O/ Mo/ M subsp. pseudoplantaginea (Skottsb.) G. D. Carr end; O (Ko) Dubautia linearis (Gaudich.) D. D. Keck subsp. hillebrandii (H. Mann) G. D. Carr end; H subsp. linearis end; L/ M Dubautia menziesii (A. Gray) D. D. Keck end; EM Dubautia microcephala Skottsb. end; K; Conservation Assessment = R Dubautia paleata A. Gray end; K Dubautia pauciflorula H. St. John & G. D. Carr end; K (Wahiawa Stream); Conservation Assessment = E; US Status = E Dubautia plantaginea Gaudich. subsp. humilis G. D. Carr end; WM (Black Gorge); Conservation Assessment = E; US Status = E subsp. magnifolia (Sherff) G. D. Carr end; K; Conservation Assessment = R; US Status = E

subsp. plantaginea end; O/ Mo/ L/ M/ H Dubautia platyphylla (A. Gray) D. D. Keck end; M; Conservation Assessment = R Dubautia raillardioides Hillebr. end; K Dubautia reticulata (Sherff) D. D. Keck end; M; Conservation Assessment = R Dubautia scabra (DC.) D. D. Keck subsp. leiophylla (A. Gray) G. D. Carr end; Mo/ L/ M/ H (Kohala Mts) subsp. scabra end; EM/ H Dubautia sherffiana Fosberg end; O (Wa); Conservation Assessment = V Dubautia syndetica G. D. Carr & Lorence end; K (Wahiawa Mts); Conservation Assessment = R Dubautia waialealae Rock end; K (Wai‘ale‘ale); Conservation Assessment = R; US Status = E Dubautia waianapanapaensis G. D. Carr end; EM Hesperomannia arborescens A. Gray end; O (Ko)/ Mo (Oloku‘i; Pelekunu)/ L (ex)/ WM (Kahakuloa, Honokōhau Valley); Conservation Assessment = E; US Status = E Hesperomannia arbuscula Hillebr. end; O (Wa)/ WM; Conservation Assessment = E; US Status = E Hesperomannia lydgatei C. N. Forbes end; K (Wahiawa Stream); Conservation Assessment = E; US Status = E Keysseria erici (C. N. Forbes) Cabrera end; K (Alaka‘i Swamp; Mt Wai‘ale‘ale; Wainiha Valley); Conservation Assessment = R; US Status = E Keysseria helenae (C. N. Forbes & Lydgate) Cabrera end; K (Alaka‘i Swamp; Mt Wai‘ale‘ale; Kilohana; Kahōluamanu); Conservation Assessment = R; US Status = E Keysseria maviensis (H. Mann) Cabrera end; south Mo (Kalapamoa Ridge)/ WM (Mt ‘Eke; Pu‘ukukui)/ EM (Kuiki; Kaupō Gap; Kīpahulu and Waiho‘i valleys); Conservation Assessment = V Lipochaeta connata (Gaudich.) DC. subsp. acris (Sherff) W. L. Wagner & H. Rob. end; Ni/ K subsp. connata end; K/ WM Lipochaeta degeneri Sherff end; southwest Mo (extinct); Conservation Assessment = EX Lipochaeta heterophylla A. Gray end; Mo/ L/ M Lipochaeta lobata (Gaudich.) DC. subsp. leptophylla (O. Deg. & Sherff) W. L. Wagner & H. Rob. end; O (Wa: Kolekole Pass; Kānehoa); Conservation Assessment = E; US Status = E subsp. lobata end; Ni/ O/ WM

APPENDIX

Lipochaeta rockii Sherff end; Mo/ M/ Ka/ H? Lipochaeta succulenta (Hook. & Arn.) DC. end; Ni/ K/ O (r)/ Mo/ M/ Ka/ H Melanthera bryanii (Sherff) W. L. Wagner & H. Rob. end; Ka (extinct); Conservation Assessment = EX Melanthera fauriei (H. Lév.) W. L. Wagner & H. Rob. end; K (Koai‘e and Olokele canyons; Wai‘alae Valley; Hikimoe and Po‘opo‘oiki valleys); Conservation Assessment = E; US Status = E Melanthera integrifolia (Nutt.) W. L. Wagner & H. Rob. end; Ku/ La/ HI Melanthera kamolensis (O. Deg. & Sherff) W. L. Wagner & H. Rob. end; southeast EM (Kamole and Kepuni Gulches); Conservation Assessment = E; US Status = E Melanthera lavarum (Gaudich.) W. L. Wagner & H. Rob. end; Mo/ L/ M/ Ka/ northwest H Melanthera micrantha (Nutt.) W. L. Wagner & H. Rob. subsp. exigua (O. Deg. & Sherff) W. L. Wagner & H. Rob. end; K (Hā‘upu Ridge); Conservation Assessment = E; US Status = E subsp. micrantha end; K (Olokele and Hanapēpē valleys); Conservation Assessment = E; US Status = E Melanthera perdita (Sherff) W. L. Wagner & H. Rob. end; Ni (Kawaihoa Pt) (extinct); Conservation Assessment = EX Melanthera populifolia (Sherff) W. L. Wagner & H. Rob. end; L (Maunalei Valley) (extinct); Conservation Assessment = EX Melanthera remyi (A. Gray) W. L. Wagner & H. Rob. end; O (northwest Wa); Conservation Assessment = R Melanthera subcordata (A. Gray) W. L. Wagner & H. Rob. end; H (North Kona District; South Kohala District) Melanthera tenuifolia (A. Gray) W. L. Wagner & H. Rob. end; O (central Wa); Conservation Assessment = E; US Status = E Melanthera tenuis (O. Deg. & Sherff) W. L. Wagner & H. Rob. end; O (central Wa); Conservation Assessment = V Melanthera venosa (Sherff) W. L. Wagner & H. Rob. end; H (Nohonaohae, Holoholokū, and Heihei cinder cones; Pu‘upā; South Kohala District); Conservation Assessment = E; US Status = E Melanthera waimeaensis (H. St. John) W. L. Wagner & H. Rob. end; K (Waimea Canyon); Conservation Assessment = E; US Status = E Pseudognaphalium sandwicensium (Gaudich.) Anderb. var. hawaiiense (O. Deg. & Sherff) W. L. Wagner end; EM (Auwahi)/ H var. kilaueanum (O. Deg. & Sherff) W. L. Wagner end; H var. molokaiense (O. Deg. & Sherff) W. L. Wagner end; O (Ko: Waimānalo, Diamond Head)/ west Mo/ L (Pu‘u Kilea)/ EM (Waiehu sandhills); Conservation Assessment = V; US Status = C var. sandwicensium end; Ku/ Mi/ Ni/ K/ O/ Mo/ L/ M/ H Remya kauaiensis Hillebr. end; K (Kōke‘e); Conservation Assessment = E; US Status = E

Remya mauiensis Hillebr. end; WM (Manawainui Gulch; Pāpalaua; formerly Olowalu Canyon; ‘Īao Valley; and behind Lahaina); Conservation Assessment = E; US Status = E Remya montgomeryi W. L. Wagner & D. R. Herbst end; K (Kalalau); Conservation Assessment = E; US Status = E Tetramolopium arenarium (A. Gray) Hillebr. subsp. arenarium end; EM (Kula, ex)/ H (Hulalālai, Nohonaohae, Waimea); Conservation Assessment = E; US Status = E subsp. laxum Lowrey end; EM (extinct); Conservation Assessment = EX; US Status = E Tetramolopium capillare (Gaudich.) H. St. John end; WM (Lahaina Luna to Wailuku; extant only at Kaua‘ula); Conservation Assessment = E; US Status = E Tetramolopium consanguineum (A. Gray) Hillebr. subsp. consanguineum end; K? (extinct); Conservation Assessment = EX subsp. leptophyllum (Sherff) Lowrey end; H (Ka‘ū District, between Mauna Loa and Mauna Kea); Conservation Assessment = R Tetramolopium conyzoides (A. Gray) Hillebr. end; southwest Mo/ L/ WM (Waiehu?)/ EM (Kula)/ H (extinct); Conservation Assessment = EX Tetramolopium filiforme Sherff var. filiforme end; O (Wa: ‘Ōhikilolo Ridge; Kea‘au Valley); Conservation Assessment = E; US Status = E var. polyphyllum (Sherff) Lowrey end; O (Wa: ‘Ōhikilolo Ridge); Conservation Assessment = E; US Status = E Tetramolopium humile (A. Gray) Hillebr. subsp. haleakalae Lowrey end; EM subsp. humile end; H (Mauna Loa; Mauna Kea; Hualālai; Kīlauea) Tetramolopium lepidotum (Less.) Sherff subsp. arbusculum (A. Gray) Lowrey end; EM (Haleakalā Crater) (extinct); Conservation Assessment = EX subsp. lepidotum end; O (Wa)/ L (extinct); Conservation Assessment = E; US Status = E Tetramolopium remyi (A. Gray) Hillebr. end; L (Awalua Ridge; Lāna‘ihale)/ WM (between Lahaina and Wailuku, ex); Conservation Assessment = E; US Status = E Tetramolopium rockii Sherff var. calcisabulorum (H. St. John) Lowrey end; northwest Mo (Mo‘omomi); Conservation Assessment = V; US Status = T var. rockii end; northwest Mo (Mo‘omomi); Conservation Assessment = V; US Status = T Tetramolopium sylvae Lowrey end; windward Mo (Ho‘olehua to Pelekunu Valley)/ WM (Kahakuloa); ConservationAssessment = R

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Tetramolopium tenerrimum (Less.) Nees end; O (Ko) (extinct); Conservation Assessment = EX Wilkesia gymnoxiphium A. Gray end; K (Waimea Canyon) Wilkesia hobdyi H. St. John end; K (Polihale; Kā‘aweiki); Conservation Assessment = E; US Status = E Begoniaceae Hillebrandia sandwicensis Oliv. end; K/ O (Wa: Mt Ka‘ala, ex)/ Mo/ M; Conservation Assessment = V Boraginaceae Cordia subcordata Lam. ind; Ni/ K/ O/ Mo (Pōhakupili Stream)/ L/ M/ H/ Heliotropium anomalum Hook. & Arn. var. argenteum A. Gray end; Ni/ K/ O/ Mo/ M (rare)/ H (rare); perhaps formerly L & Ka Heliotropium curassavicum L. ind; La/ FF/ N/ HI Nama sandwicensis A. Gray end; Li/ La/ Ni/ K/ O/ Mo/ L/ M/ H; Conservation Assessment = V Brassicaceae Lepidium arbuscula Hillebr. end; O (leeward Wa); Conservation Assessment = E; US Status = E Lepidium bidentatum Montin var. o-waihiense (Cham. & Schltdl.) Fosberg end; Ku/ Mi (ex)/ PH/ La (ex)/ K/ O/ Mo/ L/ M/ H; Conservation Assessment = V Lepidium orbiculare H. St. John end; K (Hā‘upu Ridge); Conservation Assessment = E Lepidium remyi Drake end; H (extinct); Conservation Assessment = EX Lepidium serra H. Mann end; K Rorippa sarmentosa (G. Forst. ex DC.) J. F. Macbr. ind; K/ O/ Mo/ M/ H pol? Campanulaceae Brighamia insignis A. Gray end; Ni (Ka‘ali Cliff, extinct)/ K (Nāpali Coast, Hā‘upu Ridge, Nāwiliwili); Conservation Assessment = E; US Status = E Brighamia rockii H. St. John end; windward Mo (Kalaupapa to Hālawa)/ L (Maunalei Valley, extinct)/ M (extinct); Conservation Assessment = E; US Status = E Clermontia arborescens (H. Mann) Hillebr. subsp. arborescens end; WM (‘Īao Valley and Hana‘ula); Conservation Assessment = R subsp. waihiae (Wawra) Lammers end; WM (northern)/ EM subsp. waikoluensis (H. St. John) Lammers end; Mo/ L

Clermontia calophylla E. Wimm. end; H (Kohala Mts); Conservation Assessment = R Clermontia clermontioides (Gaudich.) A. Heller subsp. clermontioides end; H (Mauna Loa and Hualālai: Pāhala to Pu‘u Lehua) subsp. rockiana (E. Wimm.) Lammers end; H (Mauna Loa and Hualālai: Kealia to Pu‘uwa‘awa‘a) Clermontia drepanomorpha Rock end; H (Kohala Mts); Conservation Assessment = E; US Status = E Clermontia fauriei H. Lév. end; K/ O (Wa: Makaleha Valley; Ko: Kōloa Gulch) Clermontia grandiflora Gaudich. subsp. grandiflora end; WM subsp. maxima Lammers end; EM; Conservation Assessment = E subsp. munroi (H. St. John) Lammers end; Mo/ L/ M Clermontia hawaiiensis (Hillebr.) Rock end; H (Puna and Ka‘ū districts) Clermontia kakeana Meyen end; O/ Mo/ M Clermontia kohalae Rock end; H (Kohala, Hāmākua, and North Hilo districts) Clermontia lindseyana Rock end; EM (leeward slope Haleakalā)/ H (South Hilo, North Hilo, Ka‘ū, and South Kona districts); Conservation Assessment = E; US Status = E Clermontia micrantha (Hillebr.) Rock end; L (Kumoa Gulch)/ WM (Pu‘ukukui; Mt ‘Eke; Lanilili) Clermontia montis-loa Rock end; H (North Hilo, South Hilo, Puna, and Ka‘ū districts) Clermontia multiflora Hillebr. end; O (Ko: Wailupe)/ WM (Waihe‘e) (extinct); Conservation Assessment = EX Clermontia oblongifolia Gaudich. subsp. brevipes (E. Wimm.) Lammers end; Mo; Conservation Assessment = E; US Status = E subsp. mauiensis (Rock) Lammers end; L (extinct)/ M; Conservation Assessment = E; US Status = E subsp. oblongifolia end; O Clermontia pallida Hillebr. end; Mo Clermontia parviflora Gaudich. ex A. Gray end; H (Kohala Mts; windward slopes of Mauna Kea and Mauna Loa) Clermontia peleana Rock subsp. peleana end; H (windward slopes of Mauna Kea and formerly Mauna Loa) (extinct?); Conservation Assessment = EX?; US Status = E subsp. singuliflora (Rock) Lammers end; EM (windward slopes of Haleakalā/ H (windward slopes of Mauna Kea) (extinct); Conservation Assessment = EX; US Status = E

APPENDIX

Clermontia persicifolia Gaudich. end; O Clermontia pyrularia Hillebr. end; H (windward Mauna Kea; leeward Mauna Loa); Conservation Assessment = E; US Status = E Clermontia samuelii C. N. Forbes subsp. hanaensis (H. St. John) Lammers end; EM (Kūhiwa Gulch to Pu‘uiki); Conservation Assessment = V; US Status = E subsp. samuelii end; EM (Kīpahulu to Wai‘ānapanapa); Conservation Assessment = V; US Status = E Clermontia tuberculata C. N. Forbes end; EM; Conservation Assessment = R Clermontia waimeae Rock end; H (Kohala Mts); Conservation Assessment = R Cyanea aculeatiflora Rock end; EM (windward slope Haleakalā) Cyanea acuminata (Gaudich.) Hillebr. end; O (Ko); Conservation Assessment = R; US Status = E Cyanea angustifolia (Cham.) Hillebr. end; O/ east Mo/ L/ north WM Cyanea arborea Hillebr. end; EM (leeward slope Haleakalā) (extinct); Conservation Assessment = EX Cyanea asarifolia H. St. John end; northeast K (Anahola Stream); Conservation Assessment = E; US Status = E Cyanea asplenifolia (H. Mann) Hillebr. end; north WM; Conservation Assessment = E; US Status = PE Cyanea calycina (Cham.) Lammers end; O (Wa; rare Ko); Conservation Assessment = R; US Status = E Cyanea comata Hillebr. end; EM (leeward slope Haleakalā) (extinct); Conservation Assessment = EX Cyanea copelandii Rock subsp. copelandii end; H (windward slope Mauna Loa) (extinct); Conservation Assessment = EX; US Status = E subsp. haleakalaensis (H. St. John) Lammers end; EM (windward slope Haleakalā); Conservation Assessment = V; US Status = E Cyanea coriacea (A. Gray) Hillebr. end; K Cyanea crispa (Gaudich.) Lammers, Givnish & Sytsma end; O (Ko); Conservation Assessment = E; US Status = E Cyanea cylindrocalyx (Rock) Lammers end; H (Waipi‘o Valley) (extinct); Conservation Assessment = EX Cyanea dolichopoda Lammers & Lorence end; K (Blue Hole, N Fork Wailua River) (extinct?); Conservation Assessment = EX?; US Status = E Cyanea dunbariae Rock end; Mo (Waihānau and Mokomoko gulches); Conservation Assessment = E; US Status = E Cyanea duvalliorum Lammers & H. Oppenh. end; EM; US Status = PE

Cyanea eleeleensis (H. St. John) Lammers end; K (Wainiha Valley) (extinct?); Conservation Assessment = EX?; US Status = E Cyanea elliptica (Rock) Lammers end; L/ M Cyanea fissa (H. Mann) Hillebr. end; K Cyanea floribunda E. Wimm. end; H (Kohala Mts; windward Mauna Kea and Mauna Loa) Cyanea gibsonii Hillebr. end; L; Conservation Assessment = E; US Status = E Cyanea giffardii Rock end; H (Glenwood) (extinct); Conservation Assessment = EX Cyanea glabra (E. Wimm.) H. St. John end; EM (windward slope Haleakalā); Conservation Assessment = E; US Status = E Cyanea grimesiana Gaudich. subsp. grimesiana end; O (Wa: Mt Ka‘ala; Ko), Mo; Conservation Assessment = E; US Status = E subsp. obatae (H. St. John) Lammers end; O (south Wa: Pu‘uhāpapa to Ka‘aikukui); Conservation Assessment = E; US Status = E Cyanea habenata (H. St. John) Lammers end; K (Limahuli Valley); Conservation Assessment = E Cyanea hamatiflora Rock subsp. carlsonii (Rock) Lammers end; H (Kona District); Conservation Assessment = E; US Status = E subsp. hamatiflora end; EM (windward slope Haleakalā); Conservation Assessment = E; US Status = E Cyanea hardyi Rock end; K Cyanea hirtella (H. Mann) Hillebr. end; K Cyanea horrida (Rock) O. Deg. & Hosaka end; EM (windward slope Haleakalā); Conservation Assessment = E; US Status = PE Cyanea humboldtiana (Gaudich.) Lammers, Givnish & Sytsma end; O (Ko); Conservation Assessment = E; US Status = E Cyanea kahiliensis (H. St. John) Lammers end; K (Mt Kāhili); Conservation Assessment = R Cyanea kauaulaensis H. Oppenheimer & Lorence end; WM (Kaua‘ula and Waikapu valleys); Conservation Assessment = E Cyanea kolekoleensis (H. St. John) Lammers end; K (Wahiawa drainage); Conservation Assessment = EX? US Status = E Cyanea koolauensis Lammers, Givnish & Sytsma end; O (Ko); Conservation Assessment = E; US Status = E Cyanea kuhihewa Lammers end; K (Limahuli Valley) (extinct?); Conservation Assessment = EX?; US Status = E Cyanea kunthiana Hillebr. end; M; Conservation Assessment = R; US Status = PE Cyanea lanceolata (Gaudich.) Lammers, Givnish & Sytsma end; O (Ko); Conservation Assessment = R; US Status = E

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Cyanea leptostegia A. Gray end; west K; Conservation Assessment = V Cyanea linearifolia Rock end; K (Wahiawa Mts) (extinct); Conservation Assessment = EX Cyanea lobata H. Mann subsp. baldwinii (C. N. Forbes & G. C. Munro) Lammers end; L (Lāna‘ihale [ex?]; Pu‘u Ali‘i); Conservation Assessment = E; US Status = E subsp. lobata end; WM (northern); Conservation Assessment = E; US Status = E Cyanea longiflora (Wawra) Lammers, Givnish & Sytsma end; O (Wai‘anae Mts; cent. Ko‘olau Mts); Conservation Assessment = E; US Status = E Cyanea longissima (Rock) H. St. John end; EM (windward slope Haleakalā) (extinct); Conservation Assessment = EX Cyanea macrostegia Hillebr. end; M Cyanea magnicalyx Lammers end; WM; Conservation Assessment = E; US Status = PE Cyanea mannii (Brigham) Hillebr. end; central Mo; Conservation Assessment = E; US Status = E Cyanea maritae Lammers & H. Oppenh. end; EM; US Status = PE Cyanea marksii Rock end; H (Kona District); Conservation Assessment = E; US Status = PE Cyanea mauiensis (Rock) Lammers end; M (extinct); Conservation Assessment = EX; US Status = PE Cyanea mceldowneyi Rock end; EM (windward slope Haleakalā); Conservation Assessment = E; US Status = E Cyanea membranacea Rock end; O (Wa); Conservation Assessment = R Cyanea minutiflora Lammers end; K (Ka Loko Reservoir) (extinct); Conservation Assessment = EX Cyanea munroi (Hosaka) Lammers end; Mo/ L; Conservation Assessment = E; US Status = PE Cyanea obtusa (A. Gray) Hillebr. end; WM/ EM (S slope of Haleakalā and Makawao Forest Reserve); Conservation Assessment = E; US Status = PE Cyanea parvifolia (C. N. Forbes) Lammers, Givnish & Sytsma end; K (Wai‘oli Valley) (extinct); Conservation Assessment = EX Cyanea pilosa A. Gray subsp. longipedunculata (Rock) Lammers end; H (windward Mauna Kea and Mauna Loa) subsp. pilosa end; H (Kohala Mts; windward Mauna Kea) Cyanea pinnatifida (Cham.) E. Wimm. end; O (central Wa); Conservation Assessment = E; US Status = E Cyanea platyphylla (A. Gray) Hillebr. end; H (Waipi‘o Valley; Puna District; windward Mauna

Kea and Mauna Loa); Conservation Assessment = E; US Status = E Cyanea pohaku Lammers end; EM (Pu‘unianiau northwest Haleakalā) (extinct); Conservation Assessment = EX Cyanea procera Hillebr. end; Mo (eastern); Conservation Assessment = E; US Status = E Cyanea profuga C. N. Forbes end; Mo (Wāwā‘ia and Kumu‘eli gulches); Conservation Assessment = E; US Status = PE Cyanea pseudofauriei Lammers end; K Cyanea purpurellifolia (Rock) Lammers, Givnish & Sytsma end; O (Ko: Kaluanui Valley); Conservation Assessment = E; US Status = E Cyanea pycnocarpa (Hillebr.) E. Wimm. end; H (Kohala Mts) (extinct); Conservation Assessment = EX Cyanea quercifolia (Hillebr.) E. Wimm. end; EM (leeward slope Haleakalā) (extinct); Conservation Assessment = EX Cyanea recta (Wawra) Hillebr. end; K (Makaleha Mts; Powerline Trail; Wai‘oli Valley); Conservation Assessment = V; US Status = T Cyanea remyi Rock end; K (Wailua, Wahiawa, and Wai‘oli valleys); Conservation Assessment = E; US Status = E Cyanea rivularis Rock end; K; Conservation Assessment = E; US Status = E Cyanea salicina H. Lév. end; K (northeastern); Conservation Assessment = E; US Status = T Cyanea scabra Hillebr. end; WM Cyanea sessilifolia (O. Deg.) Lammers end; O (N Ko‘olau Mts); Conservation Assessment = E; US Status = E Cyanea shipmanii Rock end; H (windward slope Mauna Kea); Conservation Assessment = E; US Status = E Cyanea solanacea Hillebr. end; Mo/ WM; Conservation Assessment = E; US Status = PE Cyanea solenocalyx Hillebr. end; east Mo; Conservation Assessment = R Cyanea spathulata (Hillebr.) A. Heller end; west K (‘Ōpaewela; Kahōluamanu; Kōke‘e; Kalalau) Cyanea st.-johnii (Hosaka) Lammers, Givnish & Sytsma end; O (central Ko); Conservation Assessment = E; US Status = E Cyanea stictophylla Rock end; H (Kona Coast; Ka‘ū District); Conservation Assessment = E; US Status = E Cyanea superba (Cham.) A. Gray subsp. regina (Wawra) Lammers end; O (south Ko) (extinct); Conservation Assessment = EX; US Status = E subsp. superba

APPENDIX

end; O (north Wa); Conservation Assessment = E; US Status = E Cyanea tritomantha A. Gray end; H (Waipi‘o Valley; windward Mauna Kea and Mauna Loa); Conservation Assessment = V; US Status = PE Cyanea truncata (Rock) Rock end; O (windward Ko) (extinct); Conservation Assessment = EX; US Status = E Cyanea undulata C. N. Forbes end; K (Wahiawa area); Conservation Assessment = E; US Status = E Delissea argutidentata (F. E. Wimmer) H. St. John end; H (west and southwest Hualālai, one individual extant; Pu‘ulehua and south Mauna Loa, extinct); Conservation Assessment = E; US Status = E Delissea fallax Hillebr. end; H (Hāmākua and Hilo districts) (extinct); Conservation Assessment = EX Delissea fauriei H. Lev. end; Mo; Conservation Assessment = EX Delissea kauaiensis (Lammers) Lammers end; K (Hanapēpē Falls (extinct); Ku‘ia); Conservation Assessment = E Delissea laciniata Hillebr. end; O (Ko: Wailupe) (extinct); Conservation Assessment = EX Delissea lanaiensis (Rock) Lammers end; L (extinct); Conservation Assessment = EX Delissea lauliiana Lammers end; O (Ko: Wailupe) (extinct); Conservation Assessment = EX Delissea niihauensis H. St. John end; Ni (extinct); Conservation Assessment = EX Delissea parviflora Hillebr. end; H (Kohala Mts; Mauna Kea) (extinct); Conservation Assessment = EX Delissea rhytidosperma H. Mann end; K; Conservation Assessment = E; US Status = E Delissea sinuata Hillebr. end; O (Wa) (extinct); Conservation Assessment = EX Delissea subcordata Gaudich. subsp. obtusifolia (Wawra) Lammers end; O (central Ko‘olau Mts); Conservation Assessment = EX subsp. subcordata end; O (Ko‘olau Mts); Conservation Assessment = EX Delissea takeuchii Lammers end; O (Wai‘anae Mts); Conservation Assessment = EX Delissea undulata Gaudich. end; WM (extinct); Conservation Assessment = EX Delissea waianaeensis Lammers end; O (Wai‘anae Mts); Conservation Assessment = E; US Status = E Lobelia dunbariae Rock subsp. dunbariae end; Mo (Waihānau Stream) (extinct); Conservation Assessment = EX subsp. paniculata (Rock) Lammers end; Mo (Kawela; Wai‘ale‘ia; Waikolu); Conservation Assessment = R

Lobelia gaudichaudii A. DC. end; O (central and south Ko); Conservation Assessment = V Lobelia gloria-montis Rock subsp. gloria-montis end; Mo (Pāpa‘alā Pali)/ WM (Pu‘ukukui) subsp. longibracteata (Rock) Lammers end; M (rim of ‘Īao Valley and Haleakalā) Lobelia grayana E. Wimm. end; EM (Haleakalā) Lobelia hillebrandii Rock end; Mo (Waihānau Stream)/ M Lobelia hypoleuca Hillebr. end; K/ O/ Mo/ L/ M/ H Lobelia kauaensis (A. Gray) A. Heller end; K (Wahiawa Bog; Wai‘ale‘ale); Conservation Assessment = R Lobelia koolauensis (Hosaka & Fosberg) Lammers end; O (north Ko); Conservation Assessment = E; US Status = E Lobelia monostachya (Rock) Lammers end; O (south Ko); Conservation Assessment = E; US Status = E Lobelia niihauensis H. St. John end; Ni (ex)/ K/ O (north Wa); Conservation Assessment = E; US Status = E Lobelia oahuensis Rock end; O (Ko); Conservation Assessment = E; US Status = E Lobelia remyi Rock end; O (extinct); Conservation Assessment = EX Lobelia villosa (Rock) H. St. John & Hosaka end; K (Alaka‘i Swamp; Wai‘ale‘ale); Conservation Assessment = R Lobelia wahiawa Lammers end; K; Conservation Assessment = R Lobelia yuccoides Hillebr. end; K/ O (Wa); Conservation Assessment = R Trematolobelia auriculata H. St. John end; L (extinct?); Conservation Assessment = EX? Trematolobelia grandifolia (Rock) O. Deg. end; H (Kohala Mts; between Mauna Loa and Mauna Kea; Hawai‘i Volcanoes NP); Conservation Assessment = R Trematolobelia kaalae (O. Deg.) Lammers end; O (Wai‘anae Mts) Trematolobelia kauaiensis (Rock) Skottsb. end; K Trematolobelia macrostachys (Hook. & Arn.) Zahlbr. end; O (Ko‘olau Mts) / Mo/ M Trematolobelia rockii H. St. John end; Mo (extinct?); Conservation Assessment = EX? Trematolobelia singularis H. St. John end; O (Ko‘olau Mts: Kōnāhuanui); Conservation Assessment = E; US Status = E Trematolobelia wimmeri O. Deg. & I. Deg. end; H Capparidaceae Capparis sandwichiana DC. end; Mi/ PH/ La (extinct)/ HI; Conservation Assessment =V

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Caryophyllaceae Schiedea adamantis H. St. John end; O (Diamond Head); Conservation Assessment = E; US Status = E Schiedea amplexicaulis H. Mann end; K (extinct); Conservation Assessment = EX Schiedea apokremnos H. St. John end; K (Nu‘alolo Kai, Miloli‘i, and Kalalau valleys, Nāpali Coast); Conservation Assessment = E; US Status = E Schiedea attenuata W. L. Wagner, Weller & Sakai end; K (Kalalau Valley); Conservation Assessment = E; US Status = E Schiedea diffusa A. Gray subsp. diffusa end; Mo (Kawela to Waikolu valleys)/ M; Conservation Assessment = R subsp. macraei (Sherff) W. L. Wagner & Weller end; H (Kohala Mts; ‘Ōla‘a Tract, Hawai‘i Volcanoes NP); Conservation Assessment = E; US Status = PE Schiedea globosa H. Mann end; O (southeast)/ Mo (north and northwest coasts)/ M (east and west coasts)/ H (Kohala coast); Conservation Assessment = V Schiedea haleakalensis O. Deg. & Sherff end; EM (Kaupō Gap; Hōlua near Ko‘olau Gap); Conservation Assessment = E; US Status = E Schiedea hawaiiensis Hillebr. end; H (Pōhakuloa, one individual); Conservation Assessment = E; US Status = PE Schiedea helleri Sherff end; K (Kahōluamanu, Mōhihi Stream; Wai‘alae Valley); Conservation Assessment = E; US Status = E Schiedea hookeri A. Gray end; O (north and central Wa)/ EM? (Haleakalā, extinct); Conservation Assessment = E; US Status = E Schiedea implexa (Hillebr.) Sherff end; EM (Auwahi and Kaupō Gap) (extinct); Conservation Assessment = EX Schiedea jacobii W. L. Wagner, Weller & A. C. Medeiros end; EM (Hanawi); Conservation Assessment = E; US Status = PE Schiedea kaalae Wawra end; O (Wa: Makaleha; Pahole Gulch; Mokulē‘ia; Pu‘uhāpapa; Huliwai. Ko: Punalu‘u; Kaipapa‘u); Conservation Assessment = E; US Status = E Schiedea kauaiensis H. St. John end; K (northern); Conservation Assessment = E; US Status = E Schiedea kealiae Caum & Hosaka end; O (north Wa); Conservation Assessment = E; US Status = E Schiedea laui W. L. Wagner & Weller end; Mo (Waikolu drainage); Conservation Assessment = E; US Status = PE Schiedea ligustrina Cham. & Schltdl. end; O (Wa: Pālehua; Pu‘ukaua; Keawapilau Gulch; Mokulē‘ia); Conservation Assessment = V Schiedea lychnoides Hillebr.

end; K (Alaka‘i Swamp from Ke‘ākū Cave to Kahōluamanu); Conservation Assessment = E; US Status = E Schiedea lydgatei Hillebr. end; Mo (Kamalō; Makakupa‘ia and Popohua gulches; O‘oa); Conservation Assessment = E; US Status = E Schiedea mannii H. St. John end; O (Wa: Pu‘uhāpapa; Pu‘ukawiwi; Mākua Valley; Mākua-Kea‘au Ridge; Kamaile‘unu Ridge); Conservation Assessment = V Schiedea membranacea H. St. John end; K (Mahanaloa and Ku‘ia valleys; Kalalau rim); Conservation Assessment = E; US Status = E Schiedea menziesii Hook. end; L (Maunalei Valley)/ WM (Mā‘alaea; Lahaina Luna Gulch); Conservation Assessment = V Schiedea nuttallii Hook. end; O (Wa: ‘Ēkahanui Gulch to Mokulē‘ia)/ Mo (eastern end, Kamalō, Kala‘e)/ WM; Conservation Assessment = E; US Status = E Schiedea obovata (Sherff) W. L. Wagner & Weller end; O (Wa); Conservation Assessment = E; US Status = E Schiedea pentandra W. L. Wagner & E. M. Harris end; O (Wa: Maunakapu to Mt Ka‘ala); Conservation Assessment = R Schiedea perlmanii W. L. Wagner & Weller end; K (Hā‘upu and Moloa‘a Forest Reserve); Conservation Assessment = E; US Status = E Schiedea pubescens Hillebr. end; Mo (Pelekunu to Kala‘e)/ L (Lāna‘ihale, ex)/ M (Makawao; Hāmākua; Olowalu Valley; Kā‘aNāpali); Conservation Assessment = R; US Status = C Schiedea salicaria Hillebr. end; WM (Waikapu; Kaunohua; Mā‘alaea; Olowalu; Lahaina); Conservation Assessment = V; US Status = PE Schiedea sarmentosa O. Deg. & Sherff end; Mo (below Pu‘ukolekole; ‘Ōnini, Kamalō, Kawela gulches); Conservation Assessment = E; US Status = E Schiedea spergulina A. Gray end; K (Hanapēpē; Lāwai Valley; Wahiawa; Olokele and Waimea canyons); Conservation Assessment = V; US Status = T Schiedea stellarioides H. Mann end; K (Waimea; Wai‘alae; ‘Ōpaewela; Kahōluamanu; Hā‘upu Ridge; Hanakāpī‘ai); Conservation Assessment = E; US Status = E Schiedea trinervis (H. Mann) Pax & K. Hoffm. end; O (north Wa: Mt Ka‘ala from Pu‘ukalena; Pu‘ukawiwi; and Makaha Valley); Conservation Assessment = E; US Status = E Schiedea verticillata F. Br. end; N (West Palm Valley; Devil’s Slide); Conservation Assessment = E; US Status = E Schiedea viscosa H. Mann end; K (Waimea; Nāwaimaka, Wai‘alae; Kahōluamanu; west of Alaka‘i Swamp); Conservation Assessment = E; US Status = E Silene alexandri Hillebr. end; east Mo (Pu‘ukolekole; Kamalō Gulch); Conservation Assessment = E; US Status = E

APPENDIX

Silene cryptopetala Hillebr. end; EM (Haleakalā) (extinct); Conservation Assessment = EX Silene degeneri Sherff end; EM (Ko‘olau Gap, Haleakalā) (extinct); Conservation Assessment = EX Silene hawaiiensis Sherff end; H (Kīlauea; North Kona and Hāmākua districts; Saddle Rd); Conservation Assessment = R; US Status = T Silene lanceolata A. Gray end; K (extinct)/ O (south Wa: ‘Ōhikilolo)/ Mo (Pu‘ukolekole)/ L (Maunalei Valley) (extinct)/ H (Pu‘uahi, Mauna Kea); Conservation Assessment = E; US Status = E Silene perlmanii W. L. Wagner, D. R. Herbst & Sohmer end; O (south Wa: between Palikea and Pōhākea Pass) (extinct); Conservation Assessment = EX; US Status = E Silene struthioloides A. Gray end; EM (Haleakalā)/ H (Mauna Kea; single collections at Pu‘uke‘eke‘e and Hualālai) Convolvulaceae Bonamia menziesii A. Gray end; K/ O/ Mo (extinct)/ L/ M/ H; Conservation Assessment = E; US Status = E Cressa truxillensis Kunth ind; O/ Mo/ Ka Cuscuta sandwichiana Choisy end; Ni/ K/ O/ Mo/ L/ M/ H Ipomoea imperati (Vahl) Griseb. ind; Ni/ K/ O/ Mo/ M Ipomoea indica (Burm.) Merr. ind; Ku/ Mi/ Li/ La/ N/ HI Ipomoea littoralis (L.) Blume ind; O/ Mo/ M/ H? ind? Ipomoea pes-caprae (L.) R. Br. subsp. brasiliensis (L.) Ooststr. ind; Ku (Green Island)/ Mi/ Li/ La/ FF/ N/ HI Ipomoea tuboides O. Deg. & Ooststr. end; HI Jacquemontia sandwicensis A. Gray end; HI Cucurbitaceae Sicyos albus (H. St. John) I. Telford end; H (windward slopes of Mauna Kea; Mauna Loa; and Kīlauea); Conservation Assessment = E; US Status = E Sicyos anunu (H. St. John) I. Telford end; L/ H (SW slope of Kohala Mts; NE and SW slope of Mauna Kea; southeast slope of Mauna Loa) Sicyos cucumerinus A. Gray end; K/ Mo/ M/ H; Conservation Assessment = R Sicyos erostratus H. St. John end; O (Wa: Keālia Gulch)/ Mo Sicyos herbstii (H. St. John) I. Telford end; leeward K (Makaweli to Barking Sands and Polihale)/ Mo? Sicyos hillebrandii H. St. John end; EM (Kula) (extinct); Conservation Assessment = EX Sicyos hispidus Hillebr. end; Mo/ L/ M/ H (North Kona)

Sicyos lanceoloideus (H. St. John) W. L. Wagner & D. R. Herbst end; K (Kalalau Valley; Waimea Canyon)/ O (Wa); Conservation Assessment = E Sicyos lasiocephalus Skottsb. end; Mo (Kalama‘ula)/ H (north slope Hualālai; leeward slope Kohala Mts) Sicyos macrophyllus A. Gray end; H (windward slope Kohala Mts; Mauna Kea; Mauna Loa–Mauna Kea saddle); Conservation Assessment = R; US Status = C Sicyos maximowiczii Cogn. end; Ku/ PH/ Li/ La/ Ni/ O (extinct) Sicyos pachycarpus Hook. & Arn. end; La/ N/ HI Sicyos semitonsus H. St. John end; La; Conservation Assessment = R Sicyos waimanaloensis H. St. John end; K (Waimea Canyon)/ O (extinct)/ Mo; Conservation Assessment = R Dipentodontaceae Perrottetia sandwicensis A. Gray end; K/ O/ Mo/ L/ M/ H Droseraceae Drosera anglica Huds. ind; K Ebenaceae Diospyros hillebrandii (Seem.) Fosberg end; K/ O Diospyros sandwicensis (A. DC.) Fosberg end; K/ O/ Mo/ L/ M/ H Elaeocarpaceae Elaeocarpus bifidus Hook. & Arn. end; K/ O Ericaceae Leptecophylla tameiameiae (Cham. & Schltdl.) C. M. Weiller ind; K/ O/ Mo/ L/ M/ H (perhaps on Ni and Ka in the past) Vaccinium calycinum Sm. end; K/ O/ Mo/ L/ M/ H Vaccinium dentatum Sm. end; K/ O/ Mo/ L/ M/ H Vaccinium reticulatum Sm. end; K (r)/ O (r)/ Mo (r)/ M/ H Euphorbiaceae Claoxylon sandwicense Müll. Arg. end; K/ O/ Mo (Kala‘e; r)/ L (Lāna‘ihale)/ WM (Honokōwai)/ EM (Haleakalā)/ H (Pu‘uwa‘awa‘a; Kohala; Hilo) Euphorbia arnottiana Endl. end; O (south Ko); Conservation Assessment = R Euphorbia atrococca A. Heller end; western K Euphorbia celastroides Boiss. var. amplectens Sherff end; HI

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var. celastroides end; N/ Ni/ K var. hanapepensis Sherff end; K var. kaenana Sherff end; O; Conservation Assessment = E; US Status = E var. laehiensis O. Deg., I. Deg. & Sherff end; L/ EM (Manawainui); Conservation Assessment =E var. lorifolia (A. Gray) Sherff end; L (r)/ M; Conservation Assessment = V var. stokesii (C. N. Forbes) Sherff. end; Ni/ K/ Mo/ Ka var. tomentella (Boiss.) Oudejans end; O (Wa) (extinct?); Conservation Assessment = EX? Euphorbia clusiifolia Hook. & Arn. end; O (Ko: west slopes) Euphorbia degeneri Sherff end; Ni/ K/ O/ Mo/ M/ H Euphorbia deppeana Boiss. end; O (Ko: Nu‘uanu Pali; south O); Conservation Assessment = E; US Status = E Euphorbia eleanoriae (Lorence & W. L. Wagner) Govaerts end; K (Nāpali); Conservation Assessment = R; US Status = E Euphorbia haeleeleana D. R. Herbst end; K (Ku‘ia Valley; Mahanaloa Valley; Hā‘ele‘ele Valley; Waimea Canyon)/ O (Wa: Kaluakauila Gulch); Conservation Assessment = E; US Status = E Euphorbia halemanui Sherff end; northwest K; Conservation Assessment = E; US Status = E Euphorbia herbstii (W. L. Wagner) Oudejans end; O (Wa); Conservation Assessment = E; US Status = E Euphorbia kuwaleana O. Deg. & Sherff end; O (Wa; single collections from Mokumanu and Kāne‘ohe); Conservation Assessment = E; US Status = E Euphorbia multiformis Hook. & Arn. var. microphylla Boiss. end; O/ Mo/ WM (r)/ leeward H (r) var. multiformis end; O/ M Euphorbia olowaluana Sherff end; WM/ H; Conservation Assessment = V Euphorbia remyi A. Gray ex Boiss. var. hanaleiensis Sherff end; K (Hanalei) (extinct); Conservation Assessment = EX var. kauaiensis O. Deg. & Sherff end; K (Kahōluamanu); Conservation Assessment = R; US Status = E var. remyi end; K; Conservation Assessment = R; US Status = E Euphorbia rockii C. N. Forbes end; O (Ko); Conservation Assessment = E; US Status = E Euphorbia skottsbergii Sherff var. skottsbergii end; southwest O/ northwest Mo; Conservation Assessment = R; US Status = E

var. vaccinioides Sherff end; Mo/ south M (single collection)/ Ka; Conservation Assessment = V Euphorbia sparsiflora A. Heller end; K (Wahiawa Bog); Conservation Assessment = R Fabaceae Acacia koa A. Gray end; K/ O/ Mo/ L/ M/ H Acacia koaia Hillebr. end; K/ O (Ko?)/ Mo/ L/ M/ H; Conservation Assessment = V Caesalpinia bonduc (L.) Roxb. ind; Mi (Sand Island)/ La (Pac. Remote Isl. Natl. Wildlf. Refuge)/ Ni/ K/ O/ Mo/ EM/ H Canavalia galeata (Gaudich.) Vogel end; O Canavalia hawaiiensis O. Deg., I. Deg. & J. D. Sauer end; L/ M/ H (South Pt; Hualālai; Mauna Loa; Ka‘ū Desert) Canavalia kauaiensis J. D. Sauer end; K Canavalia molokaiensis O. Deg., I. Deg. & J. D. Sauer end; east Mo; Conservation Assessment = E; US Status = E Canavalia napaliensis H. St. John end; K (Makaha to Waiahuakua valleys); Conservation Assessment = V; US Status = E Canavalia pubescens Hook. & Arn. end; Ni/ K (Nāpali Coast)/ L/ leeward EM; Conservation Assessment = V; US Status = PE Entada phaseoloides (L.) Merr. ind; K (Powerline Trail) ind? Erythrina sandwicensis O. Deg. end; HI Kanaloa kahoolawensis Lorence & K. R. Wood end; Ka (‘Ale‘ale stack); Conservation Assessment = E; US Status = E Mezoneuron kavaiense (H. Mann) Hillebr. end; K (extinct)/ O (Wa)/ L (Puhi‘elelu) (extinct?)/ WM (extinct)/ H (North Kona District); Conservation Assessment = E; US Status = E Mucuna gigantea (Willd.) DC. subsp. gigantea ind; K/ O/ Mo/ M/ H Mucuna persericea (Wilmot-Dear) T. M. Moura & A. M. G. Azebedo end; EM (Makawao to Wailua Iki); Conservation Assessment = E, US Status = PE Mucuna sloanei Fawc. & Rendle ind; K/ O/ M/ H Senna gaudichaudii (Hook. & Arn.) H. S. Irwin & Barneby ind; K/ O/ Mo/ L/ M/ Ka/ H Sesbania tomentosa Hook. & Arn. end; Ne/ N/ HI (ex on Ni‘ihau); Conservation Assessment = E; US Status = E Sophora chrysophylla (Salisb.) Seem. end; K/ O/ Mo/ L/ M/ H Strongylodon ruber Vogel end; K/ O/ Mo/ M/ H; Conservation Assessment = R Vicia menziesii Spreng. end; H (Keauhou-Kīlauea; Pu‘uwa‘awa‘a); Conservation Assessment = E; US Status = E

APPENDIX

Vigna adenantha (G. Mey.) Maréchal, Mascherpa & Stainier ind; O (Diamond Head) (extinct)/ H (extinct); Conservation Assessment = EXH Vigna marina (Burm.) Merr. ind; K/ O/ Mo/ M/ H Vigna o-wahuensis Vogel end; Ni (extinct)/ O (extinct)/ Mo/ L/ M/ Ka/ H; Conservation Assessment = E; US Status = E Gentianaceae Schenkia sebaeoides Griseb. end; K/ O (extinct)/ Mo/ L (Maunalei Canyon)/ WM; Conservation Assessment = E; US Status = E Geraniaceae Geranium arboreum A. Gray end; EM (north and west Haleakalā; Luala‘ilua Hills; south slope of Haleakalā); Conservation Assessment = E; US Status = E Geranium cuneatum Hook. subsp. cuneatum end; H (Humu‘ula; Mauna Kea; Hualālai; South Kona) subsp. hololeucum (A. Gray) Carlquist & Bissing end; H (Mauna Kea; Kahuku; ‘Āinahou; above Kūlani Prison, Mauna Loa) subsp. hypoleucum (A. Gray) Carlquist & Bissing end; H (above Kūlani Prison to Kahuku, Mauna Loa) subsp. tridens (Hillebr.) Carlquist & Bissing end; EM (Haleakalā) Geranium hanaense A. C. Medeiros & H. St. John end; EM (Hāna Forest Reserve); Conservation Assessment = V; US Status = PE Geranium hillebrandii Aedo & Muñoz Garm. end; WM (Pu‘ukukui; Mt ‘Eke); Conservation Assessment = R; US Status = PE Geranium kauaiense (Rock) H. St. John end; K (Alaka‘i Swamp to Mt Wai‘ale‘ale); Conservation Assessment = R; US Status = E Geranium multiflorum A. Gray end; EM (Haleakalā); Conservation Assessment = E; US Status = E Gesneriaceae Cyrtandra biserrata Hillebr. end; Mo (eastern: Wailau Valley; Pūko‘o Valley; Mapulehu Valley; Oloku‘i)/ EM (Kahikinui); Conservation Assessment = R Cyrtandra calpidicarpa (Rock) H. St. John & Storey end; O (Ko: Lā‘ie to Kahalu‘u on windward side; Kīpapa Gulch on leeward side) Cyrtandra confertiflora (Wawra) C. B. Clarke end; K (Nāpali Coast; Kīlauea-Anahola; Wahiawa-Hanapēpē) Cyrtandra cordifolia Gaudich. end; O (Ko: Waikāne, Kahana, and Kōnāhuanui on windward side; Halemano to Wailupe on leeward side) Cyrtandra crenata H. St. John & Storey end; O (Ko: Kahana Valley; Waikāne-Schofield Trail) (extinct); Conservation Assessment = EX; US Status = E

Cyrtandra cyaneoides Rock end; K (Makaleha and Nāmolokama Mts; Wai‘alae, Wainiha, and Wai‘oli valleys); Conservation Assessment = E; US Status = E Cyrtandra dentata H. St. John & Storey end; O (north Wa; leeward north Ko: Pa‘ala‘a; Kawailoa divide to ‘Ōpae‘ula); Conservation Assessment = E; US Status = E Cyrtandra ferripilosa H. St. John end; EM (Kīpahulu Valley, upper elev); Conservation Assessment = R; US Status = PE Cyrtandra filipes Hillebr. end; Mo (Mapulehu and Kalua‘aha valleys)/ WM (Honokōwai; Kā‘anapali; Olowalu); Conservation Assessment = R; US Status = PE Cyrtandra garnotiana Gaudich. end; O (Wa; leeward Ko: Wahiawā to Niu Valley) Cyrtandra giffardii Rock end; H (Kīlauea; Kūlani; Laupāhoehoe); Conservation Assessment = E; US Status = E Cyrtandra gracilis Hillebr. ex C. B. Clarke end; O (Ko: upper Pia Gulch; Pālolo Valley; Kōnāhuanui Gulch); Conservation Assessment = E; US Status = E Cyrtandra grandiflora Gaudich. end; O (Ko: He‘eia to Kōnāhuanui on windward side; Kīpapa Gulch to Wailupe on leeward side) Cyrtandra grayana Hillebr. end; east Mo/ L (Lāna‘ihale)/ WM Cyrtandra grayi C. B. Clarke end; Mo/ WM/ EM (Kīpahulu; Kahikinui; Pu‘u‘alaea) Cyrtandra halawensis Rock end; east Mo; Conservation Assessment = R Cyrtandra hashimotoi Rock end; M Cyrtandra hawaiensis C. B. Clarke end; O (Wa: Pu‘ukalena; Ko: Kawailoa to Wailupe Valley)/ east Mo/ M/ H Cyrtandra heinrichii H. St. John end; K (Blue Hole; Nāmolokama Mtn; upper Hanakoa; upper Hanakāpī‘ai Valley; Kekoiki); Conservation Assessment = R Cyrtandra hematos H. St. John end; Mo (Oloku‘i Plateau; Kawela; Kalua‘aha); Conservation Assessment = R Cyrtandra kalichii Wawra end; O (Wa: Mt Ka‘ala; Ko); Conservation Assessment = R Cyrtandra kamoolaensis H. St. John end; K (Kamo‘oloa Stream; ‘Ōpaeka‘a Stream); Conservation Assessment = R Cyrtandra kauaiensis Wawra end; K (Kōke‘e; Waimea drainage basin) Cyrtandra kaulantha H. St. John & Storey end; O (Ko: Waiāhole); Conservation Assessment = V; US Status = E Cyrtandra kealiae Wawra subsp. kealiae end; K; Conservation Assessment = V; US Status = T subsp. urceolata W. L. Wagner & Lorence end; K (Hanapēpē and Olokele valleys; Wahiawa Mts; Hā‘upu); Conservation Assessment = R

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Cyrtandra kohalae Rock end; H (Kohala Mts) (extinct?); Conservation Assessment = EX? Cyrtandra laxiflora H. Mann end; O (windward north Ko) Cyrtandra lessoniana Gaudich. end; O Cyrtandra longifolia (Wawra) Hillebr. ex C. B. Clarke end; K Cyrtandra lydgatei Hillebr. end; east Mo/ L (Maunalei Gulch)/ WM (Honokōhau drainage basin to Honokōwai)/EM (Honomanū Gulch); Conservation Assessment = R Cyrtandra lysiosepala (A. Gray) C. B. Clarke end; H (Kūlani; ‘Ōla‘a Tract) Cyrtandra macraei A. Gray end; O (Wa: Makaleha to Makaha valleys; Ko: Punalu‘u to Kaluanui and Kahana Iki Stream to Niu) Cyrtandra macrocalyx Hillebr. end; Mo (Waikolu to Wailau valleys; Kaunakakai Gulch)/ WM (‘Īao Valley) Cyrtandra menziesii Hook. & Arn. end; H (South Kona, Ka‘ū, and North Kona districts); Conservation Assessment = V Cyrtandra munroi C. N. Forbes end; L (Lāna‘ihale)/ WM (Makamaka‘ole); Conservation Assessment = E; US Status = E Cyrtandra nanawaleensis H. St. John end; H (low elev, Puna District); Conservation Assessment = R; US Status = PE Cyrtandra oenobarba H. Mann end; K (Kīlauea-Hā‘ena; Wahiawa-Hanapēpē; Olokele Canyon); Conservation Assessment = R; US Status = E Cyrtandra olona C. N. Forbes end; K (Wahiawa Mts) (extinct); Conservation Assessment = EX Cyrtandra oxybapha W. L. Wagner & D. R. Herbst end; WM (upper Pōhākea Gulch; Hana‘ula); Conservation Assessment = V; US Status = PE Cyrtandra paliku W. L. Wagner, K. R. Wood & Lorence end; K (Makaleha Mts); Conservation Assessment = R; US Status = E Cyrtandra paludosa Gaudich. var. microcarpa Wawra end; K var. paludosa end; K (Kōke‘e State Park; Hā‘upu Ridge; Wai‘ale‘ale)/ O (Wa: Palikea; Ko)/ M/ H Cyrtandra pickeringii A. Gray end; K (Wahiawa Mts; Hanalei; Kīlauea; Wainiha); Conservation Assessment = R Cyrtandra platyphylla A. Gray end; M/ H Cyrtandra polyantha C. B. Clarke end; O (east Ko: Niu and Kuli‘ou‘ou valleys); Conservation Assessment = E; US Status = E Cyrtandra procera Hillebr. end; Mo (Waikolu to Pelekunu valleys; Pēpē‘ōpae Bog) Cyrtandra propinqua C. N. Forbes

end; O (north Ko: Kaunala to Kīpapa and Waiāhole gulches) Cyrtandra pruinosa H. St. John & Storey end; O (Ko: Kalauao Valley) (extinct); Conservation Assessment = EX Cyrtandra rivularis H. St. John & Storey end; O (windward Ko: Kaluanui to Waikāne valleys) (extinct); Conservation Assessment = V Cyrtandra sandwicensis (H. Lév.) H. St. John & Storey end; O (leeward Ko: Pauoa to Mānoa valleys); Conservation Assessment = V Cyrtandra sessilis H. St. John & Storey end; O (windward Ko: Waikāne-Schofield Trail); Conservation Assessment = E; US Status = E Cyrtandra spathulata H. St. John end; M Cyrtandra subumbellata (Hillebr.) H. St. John & Storey end; O (north-central Ko: Kaluanui to Waiāhole valleys); Conservation Assessment = E; US Status = E Cyrtandra tintinnabula Rock end; H (Laupāhoehoe); Conservation Assessment = E; US Status = E Cyrtandra viridiflora H. St. John & Storey end; O (Ko: Lanihuli; Waikāne-Waipi‘o; Punalu‘u; Kawailoa); Conservation Assessment = E; US Status = E Cyrtandra wagneri Lorence & Perlman end; H (Laupāhoehoe NAR); Conservation Assessment = E; US Status = PE Cyrtandra waianaeensis H. St. John & Storey end; O (Wa: Mākua Valley to Palikea) Cyrtandra wainihaensis H. Lév. end; K (northeastern); Conservation Assessment = R Cyrtandra waiolani Wawra end; O (Ko: Kalihi to Kaipapa‘u valleys) (extinct); Conservation Assessment = EX; US Status = E Cyrtandra wawrae C. B. Clarke end; K Goodeniaceae Scaevola chamissoniana Gaudich. end; Mo/ L/ M/ H Scaevola coriacea Nutt. end; Ni (extinct)/ K (extinct)/ O (extinct)/ Mo (Mokuho‘oniki Islet)/ L (extinct)/ M (Waiehu; Kaupō; Mōke‘ehia Islet)/ H (extinct); Conservation Assessment = E; US Status = E Scaevola gaudichaudiana Cham. end; K/ O Scaevola gaudichaudii Hook. & Arn. end; K/ O/ Mo/ L/ M/ H Scaevola glabra Hook. & Arn. end; K/ O (Ko) Scaevola hobdyi W. L. Wagner end; WM (extinct); Conservation Assessment = EX Scaevola kilaueae O. Deg. end; H (Ocean View Estates; Ka‘ū District; Kīlauea); Conservation Assessment = R Scaevola mollis Hook. & Arn. end; K/ O/ Mo (r)

APPENDIX

Scaevola procera Hillebr. end; K/ Mo Scaevola taccada (Gaertn.) Roxb. ind; Ku/ Mi/ PH/ Li/ La/ FF/ HI Gunneraceae Gunnera kauaiensis Rock end; K (Alaka‘i Swamp; Mt Kāhili; Mt Wai‘ale‘ale) Gunnera petaloïdea Gaudich. end; K (Blue Hole)/ O/ east Mo/ M/ H (Kohala Mts) Hydrangeaceae Broussaisia arguta Gaudich. end; K/ O/ Mo/ L/ M/ H Lamiaceae Haplostachys bryanii Sherff end; central-southwest Mo (extinct); Conservation Assessment = EX Haplostachys haplostachya (A. Gray) H. St. John end; K (extinct)/ M (extinct)/ H (Kīpukakalawamauna; Pu‘ukapele [formerly from Nohonaohae]; and Waiki‘i); Conservation Assessment = E; US Status = E Haplostachys linearifolia (Drake) Sherff end; Mo (Mauna Loa; Kawela Gulch; Kamalō; Pu‘ukolekole)/ M (extinct); Conservation Assessment = EX Haplostachys munroi C. N. Forbes end; west L (Ka‘ā Ahupua‘a; Paoma‘i) (extinct); Conservation Assessment = EX Haplostachys truncata (A. Gray) Hillebr. end; M (extinct); Conservation Assessment = EX Lepechinia hastata (A. Gray) Epling ind; EM (Haleakalā) ind? Phyllostegia ambigua (A. Gray) Hillebr. end; Mo (Kapulei)/ M/ H (Mauna Loa) Phyllostegia bracteata Sherff end; M; Conservation Assessment = E; US Status = PE Phyllostegia brevidens A. Gray end; EM (Kīpahulu)/ H (Hilo; Mauna Kea); Conservation Assessment = E Phyllostegia electra C. N. Forbes end; K Phyllostegia floribunda Benth. end; H (Honokai‘a; Pāhala; Kohala Mts; Kīlauea to Laupāhoehoe); Conservation Assessment = R; US Status = PE Phyllostegia glabra (Gaudich.) Benth. var. glabra end; O (Wa: Mt Ka‘ala; Ko)/ east Mo/ L/ M var. lanaiensis Sherff end; L (known from only 2 collections) (extinct?); Conservation Assessment = EX?; US Status = E Phyllostegia grandiflora (Gaudich.) Benth. end; O (Wa: Mt Ka‘ala; Ko) Phyllostegia haliakalae Wawra end; Mo (extinct)/L (Lāna‘ihale) (extinct)/ EM (Kīpahulu Valley) ; Conservation Assessment = EX?; US Status = PE Phyllostegia helleri Sherff end; K (Wainiha Valley); Conservation Assessment = E

Phyllostegia hillebrandii H. Mann ex Hillebr. end; EM (Kula; ‘Ulupalakua) (extinct); Conservation Assessment = EX Phyllostegia hirsuta Benth. end; O (central Wa; Ko); Conservation Assessment = E; US Status = E Phyllostegia hispida Hillebr. end; Mo (east); Conservation Assessment = E; US Status = E Phyllostegia kaalaensis H. St. John end; O (Wa: Ka‘ala); Conservation Assessment = E; US Status = E Phyllostegia kahiliensis H. St. John end; K (Mt Kāhili; Nāmolokama Mtn); Conservation Assessment = R Phyllostegia knudsenii Hillebr. end; K (Koai‘e Canyon; Kōke‘e) (extinct?); Conservation Assessment = EX?; US Status = E Phyllostegia lantanoides Sherff end; O Phyllostegia macrophylla (Gaudich.) Benth. end; EM/ east H Phyllostegia mannii Sherff end; Mo (Hanalilolilo to ‘Ōhi‘alele)/ EM (‘Ukulele) (extinct); Conservation Assessment = E; US Status = E Phyllostegia micrantha H. St. John end; O (Wa: Popouwela in 1910) (extinct); Conservation Assessment = EX Phyllostegia mollis Benth. end; O (Wa; Ko: Honolulu area) (extinct); Conservation Assessment = E; US Status = E Phyllostegia parviflora (Gaudich.) Benth. var. glabriuscula A. Gray end; H (extinct); Conservation Assessment = EX; US Status = E var. lydgatei (Sherff) W. L. Wagner end; O (Wa: between Palikea and Pōhākea Pass); Conservation Assessment = E; US Status = E var. parviflora end; O/ WM (extinct); Conservation Assessment = E; US Status = E Phyllostegia pilosa H. St. John end; Mo (extinct)/ WM (above Kamalaea Bay, extinct)/ EM (Mā‘alaea, Honomanu, ‘Ulupalakua); Conservation Assessment = E; US Status = PE Phyllostegia racemosa Benth. end; H (windward slopes of Mauna Kea and Mauna Loa); Conservation Assessment = E; US Status = E Phyllostegia renovans W. L. Wagner end; K (Hanakoa; Limahuli; Wainiha); Conservation Assessment = R; US Status = E Phyllostegia rockii Sherff end; EM (‘Ukulele) (extinct); Conservation Assessment = EX Phyllostegia stachyoides A. Gray end; Mo (east)/ WM/ H (North and South Kona districts); Conservation Assessment = E Phyllostegia variabilis Bitter end; Ku/ Mi/ La (extinct); Conservation Assessment = EX

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Phyllostegia velutina (Sherff) H. St. John end; H (Kīlauea to Pu‘ukipū; Nā‘ālehu; Waipi‘o; upper Hāmākua Ditch); Conservation Assessment = E; US Status = E Phyllostegia vestita Benth. end; H (Hīlea to Laupāhoehoe; Kohala Mts) Phyllostegia waimeae Wawra end; K (Kawai‘iki); Conservation Assessment = E; US Status = E Phyllostegia warshaueri H. St. John end; H (Laupāhoehoe; Kohala Ditch Trail); Conservation Assessment = E; US Status = E Phyllostegia wawrana Sherff end; K (Hanalei; Hanakoa; Honopū; Makaleha; Kōke‘e; Wainiha); Conservation Assessment = E; US Status = E Plectranthus parviflorus Willd. ind; Ni/ K/ O/ Mo/ L/ M/ H (may also have occurred on Ka in the past) Stenogyne angustifolia A. Gray end; Mo (extinct)/ M (extinct)/ H (Pōhakuloa Training Area); Conservation Assessment = E; US Status = E Stenogyne bifida Hillebr. end; Mo (Laianui; Pelekunu Trail); Conservation Assessment = E; US Status = E Stenogyne calaminthoides A. Gray end; H Stenogyne calycosa Sherff end; M; Conservation Assessment = V Stenogyne campanulata Weller & Sakai end; K (Kalalau); Conservation Assessment = E; US Status = E Stenogyne cinerea Hillebr. end; EM (Kula) (extinct); Conservation Assessment = EX Stenogyne cranwelliae Sherff end; H (Kohala Mts); Conservation Assessment = E; US Status = PE Stenogyne haliakalae Wawra end; EM (south slope Haleakalā) (extinct); Conservation Assessment = EX Stenogyne kaalae Wawra subsp. kaalae end; O (Wa; Ko: single collection from Nu‘uanu Pali) subsp. sherffii (O. Deg.) W. L. Wagner & Weller end; O (Ko: Pe‘ahināi‘a Trail); Conservation Assessment = E Stenogyne kamehamehae Wawra end; Mo/ M Stenogyne kanehoana O. Deg. & Sherff end; O (Wa: Pu‘ukānehoa); Conservation Assessment = E; US Status = E Stenogyne kauaulaensis K. R. Wood & H. Oppenheimer end; M; Conservation Assessment = E; US Status = PE Stenogyne kealiae Wawra end; K (Kalalau to Awa‘awapuhi); Conservation Assessment = R; US Status = E Stenogyne macrantha Benth. end; H; Conservation Assessment = R Stenogyne microphylla Benth. end; EM/ H

Stenogyne oxygona O. Deg. & Sherff end; H (Kohala Mts) (extinct?); Conservation Assessment = EX? Stenogyne purpurea H. Mann end; K Stenogyne rotundifolia A. Gray end; EM (north slope Haleakalā) Stenogyne rugosa Benth. end; EM/ H Stenogyne scrophularioides Benth. end; H (Mauna Loa; Mauna Kea) Stenogyne sessilis Benth. end; L (extinct)/ M/ H Stenogyne viridis Hillebr. end; WM (Kā‘anapali) (extinct); Conservation Assessment = EX Vitex rotundifolia L. f. ind; Ni/ K/ O/ Mo/ L/ M/ H Lauraceae Cassytha filiformis L. ind; Ku/ Ni/ K/ O/ Mo/ L/ M/ H Cryptocarya mannii Hillebr. end; K/ O Loganiaceae Labordia cyrtandrae (Baill.) H. St. John end; O (Wa: Makaha Valley; Hale‘au‘au Gulch; Ko); Conservation Assessment = E; US Status = E Labordia degeneri Sherff end; K (Kōke‘e; Pihea Trail) Labordia fagraeoidea Gaudich. end; O (Ko) Labordia hedyosmifolia Baill. end; Mo/ L/ M/ H Labordia helleri Sherff end; K (Nāpali Coast); Conservation Assessment = R; US Status = E Labordia hirtella H. Mann end; east K/ O (Ko: Nu‘uanu to Wailupe valleys)/ Mo/ L/ M/ H Labordia hosakana (Sherff) W. L. Wagner, D. R. Herbst & Sohmer end; O (Ko: Kīpapa to Wa‘ahila); Conservation Assessment =R Labordia kaalae C. N. Forbes end; O (Wa); Conservation Assessment = R Labordia lorenciana K. R. Wood, W. L. Wagner & T. Motley end; K (Kawai‘iki Valley); Conservation Assessment = E Labordia lydgatei C. N. Forbes end; K (Wahiawa Mts); Conservation Assessment = E; US Status = E Labordia pumila (Hillebr.) Skottsb. end; K (Mt Wai‘ale‘ale; Alaka‘i Swamp); Conservation Assessment = R; US Status = E Labordia sessilis A. Gray end; O (Ko) Labordia tinifolia A. Gray var. lanaiensis Sherff end; L; Conservation Assessment = V; US Status = E

APPENDIX

var. tinifolia end; K/ O (Wa: Makaha Valley; Pu‘upane; Ko)/ Mo/ M/ H var. wahiawaensis H. St. John end; K (Wahiawa Valley); Conservation Assessment = E; US Status = E Labordia triflora Hillebr. end; Mo (Mapulehu Valley; Kua and Wāwā‘ia); Conservation Assessment = E; US Status = E Labordia venosa Sherff end; EM Labordia waialealae Wawra end; K (Mt Wai‘ale‘ale; Alaka‘i Swamp; Pihea; Makaleha Mts)/ possibly Mo (Kamalō) Labordia waiolani Wawra end; O/ Mo/ L (r) Malvaceae Abutilon eremitopetalum Caum end; east L (Kalulu and Maunalei valleys; Kānepu‘u; Kehewai Gulch); Conservation Assessment = E; US Status =E Abutilon incanum (Link) Sweet ind; Ni/ K/ O/ Mo/ L/ M/ Ka ind? Abutilon menziesii Seem. end; O?/ L/ M/ H; Conservation Assessment = E; US Status =E Abutilon sandwicense (O. Deg.) Christoph. end; O (Wa: between Makaleha Valley and Pu‘ukaua); Conservation Assessment = E; US Status = E Gossypium tomentosum Nutt. ex Seem. end; Ni/ K/ O/ Mo/ L/ M/ Ka; Conservation Assessment = V Hibiscadelphus bombycinus C. N. Forbes end; H (Kawaihae) (extinct); Conservation Assessment = EX Hibiscadelphus crucibracteatus Hobdy end; L (Puhi‘elelu Ridge) (extinct); Conservation Assessment = EX Hibiscadelphus distans L. E. Bishop & D. R. Herbst end; K (Koai‘e Stream; Waimea Canyon); Conservation Assessment = E; US Status = E Hibiscadelphus giffardianus Rock end; H (Kīpukapuaulu) (extinct); Conservation Assessment = EX; US Status = E Hibiscadelphus hualalaiensis Rock end; H (Hualālai and Waihou, North Kona) (extinct); Conservation Assessment = EX; US Status = E Hibiscadelphus wilderianus Rock end; EM (Auwahi, south slope Haleakalā) (extinct); Conservation Assessment = EX Hibiscadelphus woodii Lorence & W. L. Wagner end; K (Kalalau rim) (extinct?); Conservation Assessment = EX?; US Status = E Hibiscus arnottianus A. Gray subsp. arnottianus end; O (Wa; east Ko: Wahiawā to Niu Valley) subsp. immaculatus (M. J. Roe) D. M. Bates end; Mo (Wailau, Waihānau, and Pāpalaua valleys); Conservation Assessment = E; US Status = E

subsp. punaluuensis (Skottsb.) D. M. Bates end; O (Ko: Kaipapa‘u to Waiāhole) Hibiscus brackenridgei A. Gray subsp. brackenridgei end; L/ M/ H; Conservation Assessment = E; US Status = E subsp. mokuleianus (M. J. Roe) D. M. Bates end; K (Līhu‘e; Olokele Canyon) (ex)/ O (Wa: Kawaihāpai to Pu‘upane); Conservation Assessment = E; US Status = E subsp. molokaiana (Rock ex Caum) F. D. Wilson end; Mo (extinct); Conservation Assessment = EX; US Status = E Hibiscus clayi O. Deg. & I. Deg. end; east K (Nounou Mtn; Hāli‘i Valley; Anahola Mts); Conservation Assessment = E; US Status = E Hibiscus furcellatus Desr. ind; K/ O/ M/ H Hibiscus kokio Hillebr. ex Wawra subsp. kokio end; K/ O/ Mo/ M/ H?; Conservation Assessment = R subsp. saintjohnianus (M. J. Roe) D. M. Bates end; northwest K; Conservation Assessment = R Hibiscus waimeae A. Heller subsp. hannerae (O. Deg. & I. Deg.) D. M. Bates end; northwest K (Hanakāpī‘ai; Limahuli; Kalihi Wai); Conservation Assessment = E; US Status = E subsp. waimeae end; K (Waimea Canyon; west and southwest valleys) Kokia cookei O. Deg. end; west Mo (Mauna Loa) (extinct); Conservation Assessment = EX; US Status = E Kokia drynarioides (Seem.) Lewton end; H (Pu‘uwa‘awa‘a and Hu‘ehu‘e in North Kona); Conservation Assessment = E; US Status = E Kokia kauaiensis (Rock) O. Deg. & Duvel end; west K (Pa‘aiki, Ku‘ia, Mahanaloa, Kalalau, and Koai‘e valleys); Conservation Assessment = E; US Status = E Kokia lanceolata Lewton end; southeast O (Makaku; Koko Head; Wailupe Valley) (extinct); Conservation Assessment = EX Sida fallax Walp. ind; Mi/ N/ HI Thespesia populnea (L.) Sol. ex Corrêa ind; HI (except Ka) ind? Waltheria indica L. ind; Mi/ HI ind? Menispermaceae Cocculus orbiculatus (L.) DC. ind; Ni/ K/ O/ Mo/ L/ M/ H Moraceae Streblus pendulinus (Endl.) F. Muell. ind; K/ O/ Mo/ L/ M/ H Myrtaceae Eugenia koolauensis O. Deg. end; O (north Ko: Pūpūkea to Kaipapa‘u)/ Mo (Mauna Loa; extinct); Conservation Assessment = E; US Status = E

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Eugenia reinwardtiana (Blume) DC. ind; K/ O/ Mo/ M Metrosideros macropus Hook. & Arn. end; O (Ko) Metrosideros polymorpha Gaudich. var. dieteri J. W. Dawson & Stemmerm. end; K var. glaberrima (H. Lév.) H. St. John end; K/ O/ Mo/ L/ M/ H var. incana (H. Lév.) H. St. John end; K/ O/ Mo/ L/ M/ H var. macrophylla (Rock) H. St. John end; M/ H var. newellii (Rock) H. St. John end; H var. polymorpha end; O/ Mo/ L/ M/ H var. pseudorugosa (Skottsb.) J. W. Dawson & Stemmerm. end; WM var. pumila (A. Heller) J. W. Dawson & Stemmerm. end; K/ O (Ko)/ Mo/ M Metrosideros rugosa A. Gray end; O (Ko) Metrosideros tremuloides (A. Heller) Knuth end; O Metrosideros waialealae (Rock) Rock var. fauriei (H. Lév) J. W. Dawson & Stemmerm. end; Mo/ L/ WM var. waialealae end; K Syzygium sandwicense (A. Gray) Mull. Berol. end; K/ O/ Mo/ L/ M Nyctaginaceae Boerhavia acutifolia (Choisy) J. W. Moore ind; PH/ La/ HI Boerhavia herbstii Fosberg end; PH/ Li/ O/ L/ M/ Ka/ H Boerhavia repens L. ind; Ku/ Mi/ PH/ Li/ La/ FF/ N/ HI Pisonia brunoniana Endl. ind; O/ Mo/ L/ M/ H Pisonia grandis R. Br. ind; Li/ M? Pisonia sandwicensis Hillebr. end; K/ O/ Mo/ L/ M/ H Pisonia umbellifera (G. Forst.) Seem. ind; K/ O/ Mo/ L/ M/ H Pisonia wagneriana Fosberg end; K (Powerline Trail to Maunahina-Wainiha drainage); Conservation Assessment = R Oleaceae Nestegis sandwicensis (A. Gray) O. Deg., I. Deg. & L. A. S. Johnson end; K/ O/ Mo/ L/ M/ H Onagraceae Ludwigia octovalvis (Jacq.) Raven ind; K/ O/ Mo/ L/ M/ H pol?

Oxalidaceae Oxalis corniculata L. ind; Ku (Green Island)/ Mi/ HI pol? Papaveraceae Argemone glauca (Nutt. ex Prain) Pope var. decipiens Ownbey end; H (leeward side and between Mauna Loa and Mauna Kea) var. glauca end; HI (H, South Pt) Pentaphylacaceae Eurya sandwicensis A. Gray end; K/ O/ Mo/ M/ H; Conservation Assessment = R Phyllanthaceae Antidesma platyphyllum H. Mann var. hillebrandii Pax & K. Hoffm. end; K var. platyphyllum end; K/ O/ Mo/ L/ M/ H Antidesma pulvinatum Hillebr. end; O/ Mo/ L (Maunalei Gulch)/ M/ H Flueggea neowawraea W. J. Hayden end; northwest K/ O (Wa)/ Mo (extinct)/ EM (southwest slope Haleakalā)/ H (Kona Coast); Conservation Assessment = E; US Status = E Phyllanthus distichus Hook. & Arn. end; K/ O/ Mo/ L/ WM/ EM (r) Phytolaccaceae Phytolacca sandwicensis Endl. end; K/ O/ Mo/ M/ H Piperaceae Peperomia alternifolia Yunck. end; Mo/ L/ M Peperomia blanda (Jacq.) Kunth var. floribunda (Miq.) H. Huber ind; Ni/ K/ O/ Mo/ L/ M/ H Peperomia cookiana C. DC. end; K/ Mo/ M/ H Peperomia degeneri Yunck. end; Mo (Kalua‘aha Valley) (extinct?); Conservation Assessment = EX? Peperomia eekana C. DC. end; Mo/ M Peperomia ellipticibacca C. DC. end; O (Ko) Peperomia expallescens C. DC. end; Mo/ M Peperomia globulanthera C. DC. end; M Peperomia hesperomannii Wawra end; K Peperomia hirtipetiola C. DC. end; K/ L (r)/ M

APPENDIX

Peperomia hypoleuca Miq. end; H Peperomia kipahuluensis H. St. John & Lamoureux end; EM (Kīpahulu; Waiho‘i; and Kūhiwa valleys) Peperomia kokeana Yunck. end; K (Kōke‘e) Peperomia latifolia Miq. end; K/ O/ Mo/ L/ M/ H Peperomia ligustrina Hillebr. end; Mo/ M/ H Peperomia macraeana C. DC. end; O/ Mo/ L/ M/ H Peperomia mauiensis Wawra end; Mo/ L/ WM Peperomia membranacea Hook. & Arn. end; K/ O/ Mo/ M/ H Peperomia oahuensis C. DC. end; K (Hā‘upu Ridge; Kamo‘oloa Stream)/ O (Ko) Peperomia obovatilimba C. DC. end; EM/ H (Kohala Mts) Peperomia remyi C. DC. end; K/ O/ Mo/ L/ M/ H Peperomia rockii C. DC. end; east Mo; Conservation Assessment = R Peperomia sandwicensis Miq. end; K/ O/ Mo Peperomia subpetiolata Yunck. end; EM (Kula Pipeline, lower Waikamoi) (extinct?); Conservation Assessment = EX?; US Status = PE Peperomia tetraphylla (G. Forst.) Hook. & Arn. ind; K/ O/ Mo/ L/ M/ H Pittosporaceae Pittosporum argentifolium Sherff end; Mo/ M; Conservation Assessment = R Pittosporum confertiflorum A. Gray end; O/ L/ M/ H Pittosporum flocculosum (Hillebr.) Sherff end; O (Wa; Ko: Kuli‘ou‘ou–Waimānalo Ridge to Hālawa Valley) Pittosporum gayanum Rock end; K Pittosporum glabrum Hook. & Arn. end; K/ O/ Mo/ L/ M Pittosporum halophilum Rock end; Mo (Kalawao; Huelo Islet); Conservation Assessment = R; US Status = PE Pittosporum hawaiiense Hillebr. end; leeward H (Kohala Mts to Ka‘ū District); Conservation Assessment = R; US Status = PE Pittosporum hosmeri Rock end; H (Kohala Mts south to Mauna‘anu; Ka‘ū Forest Reserve) Pittosporum kauaiense Hillebr. end; K Pittosporum napaliense Sherff end; northwest K (Limahuli to Hanakoa valleys); Conservation Assessment = R; US Status = E Pittosporum terminalioides Planch. ex A. Gray end; L/ M/ leeward H (Kīlauea to Kohala Mts)

Plantaginaceae Bacopa monnieri (L.) Pennell ind; Mi/ Ni/ K/ O/ Mo/ L/ M/ H Plantago hawaiensis (A. Gray) Pilg. end; H; Conservation Assessment = E; US Status = E Plantago pachyphylla A. Gray end; K (Alaka‘i Swamp; Mt Wai‘ale‘ale)/ O (Ko)/ Mo (Pēpē‘ōpae Bog)/ M/ H (Kohala Mts; Kahuku Ranch) Plantago princeps Cham. & Schltdl. var. anomala Rock end; K (Hanapēpē and Kalalau valleys)/ O (Napepeiauolelo Gulch); Conservation Assessment = E; US Status = E var. laxifolia A. Gray end; Mo/ M/ H (extinct); Conservation Assessment = E; US Status = E var. longibracteata H. Mann end; K (Hanalei; Hanapēpē; Wahiawa Mts)/ O (Wa: Mt Ka‘ala; Ko: Kaipapa‘u to Pa‘ala‘a-Wahiawā); Conservation Assessment = E; US Status = E var. princeps end; O (Wa: scattered throughout; Ko: Mānoa Cliffs Trail; Kalihi and Nu‘uanu valleys); Conservation Assessment = E; US Status = E Plumbaginaceae Plumbago zeylanica L. ind; HI Polygonaceae Rumex albescens Hillebr. end; N/ K/ O (Wa) Rumex giganteus W. T. Aiton end; Mo/ M/ H Rumex skottsbergii O. Deg. & I. Deg. end; H (North and South Kona, Puna, and southern Ka‘ū districts) Portulacaceae Portulaca lutea Sol. ex G. Forst. ind; Ku/ Mi/ Li/ La/ GP/ FF/ Ne/ N/ HI (except Ka) Portulaca molokiniensis Hobdy end; Ml/ Ka (Kamōhio Bay; Pu‘ukoa‘e Islet); Conservation Assessment = R Portulaca sclerocarpa A. Gray end; L (Po‘opo‘o Islet)/ H; Conservation Assessment = E; US Status = E Portulaca villosa Cham. end; N/ Kl/ O/ Mo/ L/ M/ Ka/ H; Conservation Assessment =V Primulaceae Embelia pacifica Hillebr. end; K/ O/ Mo/ L/ M/ H Lysimachia daphnoides (A. Gray) Hillebr. end; K (Alaka‘i Swamp; Wahiawa Bog); Conservation Assessment = R; US Status = E Lysimachia filifolia C. N. Forbes & Lydgate

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end; K (Olokele Valley, Waikoko)/ O (Ko: Waiāhole Ditch Trail); Conservation Assessment = E; US Status = E Lysimachia forbesii Rock end; O (Ko: Castle Trail) (extinct); Conservation Assessment = EX Lysimachia glutinosa Rock end; K (Kōke‘e) Lysimachia hillebrandii Hook. f. ex A. Gray end; K (Hā‘upu to Limahuli)/ O/ east Mo/ L/ M Lysimachia iniki K. L. Marr end; K (headwaters of N Fork of Wailua River); Conservation Assessment = R; US Status = E Lysimachia kalalauensis Skottsb. end; K (Kōke‘e, from Kalalau to Makaha valleys; Hanapēpē Valley) Lysimachia lydgatei Hillebr. end; WM (Līhau; Lahaina); Conservation Assessment = E; US Status = E Lysimachia mauritiana Lam. ind; Ni/ K/ Mo/ M/ northwest H (scattered localities) Lysimachia maxima (R. Knuth) H. St. John end; east Mo (Pelekunu Trail); Conservation Assessment = E; US Status = E Lysimachia pendens K. L. Marr end; K (Wailua River; SE side of Mt Wai‘ale‘ale); Conservation Assessment = R; US Status = E Lysimachia remyi Hillebr. end; Mo/ M Lysimachia scopulensis K. L. Marr end; K (Kalalau); Conservation Assessment = R; US Status = E Lysimachia venosa (Wawra) H. St. John end; K (Mt Wai‘ale‘ale) (extinct?); Conservation Assessment = EX?; US Status = E Myrsine alyxifolia Hosaka end; K (Kōke‘e; Hā‘upu Ridge) Myrsine degeneri Hosaka end; O (Ko: Pu‘ukeahiakahoe; Kawai Nui) Myrsine denticulata (Wawra) Hosaka end; K (Alaka‘i Swamp to Mt Wai‘ale‘ale) Myrsine fernseei (Mez) Hosaka end; K (Wahiawa Bog; Powerline Trail; Hanalei River; Kaloko Reservoir) Myrsine fosbergii Hosaka end; K/ O (Ko: southeast end to Castle Trail); Conservation Assessment = R; US Status = C Myrsine helleri (O. Deg. & I. Deg.) H. St. John end; K (Wahiawa Bog; Alaka‘i Swamp to Mt Wai‘ale‘ale) Myrsine juddii Hosaka end; O (Ko: Kawai Iki and Poamoho gulches; Pe‘ahināi‘a Trail; Punalu‘u Trail); Conservation Assessment = E; US Status = E Myrsine kauaiensis Hillebr. end; K (Kōke‘e; Alaka‘i Swamp; Hanapēpē Valley) Myrsine knudsenii (Rock) Hosaka end; K (Kōke‘e; Hanapēpē Valley); Conservation Assessment = R; US Status = E Myrsine lanaiensis Hillebr. end; K/ O/ Mo/ L/ M/ H

Myrsine lessertiana A. DC. end; K/ O/ Mo/ L/ M/ H Myrsine linearifolia Hosaka end; K (west half to Pu‘uokila); Conservation Assessment = V; US Status = T Myrsine mezii Hosaka end; K (Hanapēpē); Conservation Assessment = E; US Status = E Myrsine petiolata Hosaka end; K (east half; Alaka‘i Swamp Trail) Myrsine pukooensis (H. Lév.) Hosaka end; O/ east Mo/ L/ WM Myrsine punctata (H. Lév.) Wilbur end; K (Kōke‘e Plateau; Alaka‘i Swamp; Mt Kāhili)/ O (Wa: Kuaokalā) Myrsine sandwicensis A. DC. end; O/ Mo/ L/ M/ H Myrsine vaccinioides W. L. Wagner, D. R. Herbst & Sohmer end; WM (Violet Lake, Pu‘ukukui); Conservation Assessment = E; US Status = PE Myrsine wawraea (Mez) Hosaka end; K (Kōke‘e; Alaka‘i Swamp; Mt Kāhili; Kalalau Trail) Ranunculaceae Ranunculus hawaiensis A. Gray end; EM/ H; Conservation Assessment = E; US Status = C Ranunculus mauiensis A. Gray end; K/ O (Wa: Mt Ka‘ala, ex)/ Mo (extinct)/ M/ H (extinct); Conservation Assessment = E; US Status = C Rhamnaceae Alphitonia ponderosa Hillebr. end; K/ O (r)/ Mo (r)/ L (r)/ M (r)/ H (r); Conservation Assessment = R Colubrina asiatica (L.) Brongn. ind; Ni/ K/ O/ Mo/ WM (Launiupoko) Colubrina oppositifolia Brongn. ex H. Mann end; O (Wa)/ WM (Honokōwai)/ leeward H; Conservation Assessment = E; US Status = E Gouania hillebrandii Oliv. end; Mo (Kamalō; Waiakuilani Gulch)/ L (extinct)/ WM (Pa‘upa‘u and Līhau Mts, Lahaina District)/ Ka (extinct)/ H; Conservation Assessment = E; US Status = E Gouania meyenii Steud. end; K (Hanalei and Waimea districts)/ O (Wa: Makaha– Wai‘anae Kai; Līhu‘e); Conservation Assessment = E; US Status = E Gouania vitifolia A. Gray end; O (Wa)/ WM (Lahaina, ex)/ H (Ka‘ū); Conservation Assessment = E; US Status = E Rosaceae Acaena exigua A. Gray end; K (Mt Wai‘ale‘ale, ex) (extinct?)/ WM (Pu‘ukukui) (extinct?); Conservation Assessment = EX?; US Status = E Fragaria chiloensis (L.) Duchesne subsp. sandwicensis (Decne.) Staudt end; EM/ H; Conservation Assessment = V

APPENDIX

Osteomeles anthyllidifolia (Sm.) Lindl. ind; K/ O/ Mo/ L/ M/ H Rubus hawaiensis A. Gray end; K/ Mo/ M/ H Rubus macraei A. Gray end; EM/ H; Conservation Assessment = R Rubiaceae Bobea brevipes A. Gray end; K/ O Bobea elatior Gaudich. end; K (Nāpali Coast southeast to Hā‘upu Ridge)/ O (Ko)/ Mo/ L/ M/ H (Kohala Mts) Bobea sandwicensis (A. Gray) Hillebr. end; O (Wa; south Ko: Wailupe Valley)/ Mo/ L/ M; Conservation Assessment = V Bobea timonioides (Hook. f.) Hillebr. end; K (Hā‘upu Ridge, Hōkūnui, Waimea District)/ O (Wa: Mākua; Ko: Waimano)/ EM (S Haleakalā)/ H (leeward); Conservation Assessment = V Coprosma cymosa Hillebr. end; leeward H Coprosma elliptica W. R. B. Oliv. end; K (Alaka‘i Swamp to Mt Wai‘ale‘ale; Kahōluamanu; Kōke‘e) Coprosma ernodeoides A. Gray end; WM (Mt ‘Eke?; East Bog)/ EM/ H Coprosma foliosa A. Gray end; K (r)/ O/ Mo/ L/ M Coprosma kauensis (A. Gray) A. Heller end; K Coprosma longifolia A. Gray end; O Coprosma menziesii A. Gray end; H Coprosma montana Hillebr. end; Mo (Pu‘u Ali‘i NAR)/ EM/ H Coprosma ochracea W. R. B. Oliv. end; O (Wa: Mt Ka‘ala)/ east Mo/ L/ M/ H Coprosma pubens A. Gray end; Mo/ L/ M/ H Coprosma rhynchocarpa A. Gray end; EM (Auwahi District)/ H Coprosma ternata W. R. B. Oliv. end; Mo (eastern)/ WM (Hana‘ula) Coprosma waimeae Wawra end; K Gardenia brighamii H. Mann end; O (Wa: Pu‘uku‘ua; Nānākuli)/ Mo (Mauna Loa; Mahana Flats, extinct)/ L (Kānepu‘u)/ WM (Olowalu, extinct)/ H (Pu‘uwa‘awa‘a, extinct); Conservation Assessment = E; US Status = E Gardenia mannii H. St. John & Kuykendall end; O; Conservation Assessment = E; US Status = E Gardenia remyi H. Mann end; K/ Mo/ M/ H (Hilo and Puna districts); Conservation Assessment = R; US Status = C Gynochthodes trimera (Hillebr.) Razafim & B. Bremer end; K (Hanalei District) / O (Wa; Ko: Kīpapa Gulch)/ L/ M

Kadua acuminata Cham. & Schltdl. end; K/ O/ Mo/ L/ M/ H Kadua affinis DC end; K/ O/ Mo/ L/ M/ H Kadua axillaris (Wawra) W. L. Wagner & Lorence end; Mo/ M/ H Kadua centranthoides Hook. & Arn. end; K/ O/ Mo/ L/ M/ H Kadua cookiana Cham. & Schltdl. end; K (Waiahuakua Valley; Hanakoa)/ H (Kealakekua, extinct); Conservation Assessment = E; US Status = E Kadua cordata Cham. & Schtdl. subsp. cordata end; K/ O/ Mo/ M subsp. remyi (Hillebr.) W. L. Wagner & Lorence end; L; Conservation Assessment = V; US Status = E subsp. waimeae (Wawra) W. L. Wagner & Lorence end; K Kadua coriacea (J. E. Smith) W. L. Wagner & Lorence end; O (Wa; Nu‘uanu, extinct)/ M/ H (1859 Lava Flow; Pōhakuloa); Conservation Assessment = E; US Status = E Kadua degeneri (Fosberg) W. L. Wagner & Lorence subsp. coprosmifolia (Fosberg) W. L. Wagner & Lorence end; O (Wa: Kamaile‘unu Ridge and Mokulē‘ia, north and northwest slope Mt Ka‘ala) (extinct?); Conservation Assessment = EX?; US Status = E subsp. degeneri end; O (Wa: Ka‘a‘awa and Pu‘uiki, north slope Mt Ka‘ala); Conservation Assessment = E; US Status = E Kadua elatior (H. Mann) A. Heller end; K/ O (Ko: upper Wahiawā)/ Mo/ EM Kadua fluviatilis C. N. Forbes end; K/ O (Ko: Pūpūkea to Mānoa); Conservation Assessment = R; US Status = C Kadua flynnii (W. L. Wagner & Lorence) W. L. Wagner & Lorence end; K (cliffs in northern valleys); Conservation Assessment = R Kadua foggiana (Fosberg) W. L. Wagner & Lorence end; K (Mt Kāhili; Hanalei Valley; Kōke‘e) Kadua foliosa Hillebr. end; EM (southwest slope Haleakalā) (extinct); Conservation Assessment = EX Kadua formosa Hillebr. end; WM; Conservation Assessment = R Kadua fosbergii (W. L. Wagner & D. R. Herbst) W. L. Wagner & Lorence end; O (Ko: Castle Trail to Pālolo)/ L (Lāna‘ihale) Kadua haupuensis Lorence & W. L. Wagner end; K (Hā‘upu); Conservation Assessment = EX? Kadua knudsenii Hillebr. end; K (Kōke‘e) Kadua laxiflora H. Mann end; Mo (Kawela Gulch; Mapulehu Valley)/ L/ WM; Conservation Assessment = E; US Status = E Kadua littoralis Hillebr. end; K (Princeville)/ O (extinct)/ Mo (windward)/ EM (Ke‘anae-Wailua; Hanawī; ‘Ālau Island)/ H (Honopue Valley); Conservation Assessment = V

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Kadua parvula A. Gray end; O (Wa: Pālehua to Makaleha); Conservation Assessment = E; US Status = E Kadua st.-johnii (B. C. Stone & Lane) W. L. Wagner & Lorence end; K (Honopū, Miloli‘i, Kalalau, and Nu‘alolo valleys, Nāpali Coast); Conservation Assessment = E; US Status = E Kadua tryblium (D. R. Herbst & W. L. Wagner) W. L. Wagner & Lorence end; K (Mt Kāhili; Wahiawa Stream; rim of Kalalau Valley); Conservation Assessment = R Nertera granadensis (Mutis ex L. f.) Druce ind; K/ O (Wa)/ Mo/ M / H Psychotria fauriei (H. Lév.) Fosberg end; O (Ko) Psychotria grandiflora H. Mann end; K (Kōke‘e; Alaka‘i Swamp); Conservation Assessment = E; US Status = E Psychotria greenwelliae Fosberg end; K (Kōke‘e) Psychotria hathewayi Fosberg var. brevipetiolata Fosberg end; O (Wa) var. hathewayi end; O (Wa) Psychotria hawaiiensis (A. Gray) Fosberg var. hawaiiensis end; Mo/ M/ H var. hillebrandii (Rock) Fosberg end; Mo/ M/ H var. scoriacea (Rock) Fosberg end; M/ H Psychotria hexandra H. Mann var. hexandra end; K var. oahuensis O. Deg. & Fosberg end; O (Ko); Conservation Assessment = R; US Status = E Psychotria hobdyi Sohmer end; K (Miloli‘i-Kopiwai); Conservation Assessment = E; US Status = E Psychotria kaduana (Cham. & Schltdl.) Fosberg end; K/ O/ Mo/ L/ M Psychotria mariniana (Cham. & Schltdl.) Fosberg end; K/ O/ Mo/ L/ M Psychotria mauiensis Fosberg end; K/ Mo/ L/ M/ H Psychotria wawrae Sohmer end; east K; Conservation Assessment = R Psydrax odorata (G. Forst.) A. C. Sm. & S. P. Darwin ind; K/ O/ Mo/ L/ M/ H Rutaceae Melicope adscendens (H. St. John & E. P. Hume) T. G. Hartley & B. C. Stone end; EM (Auwahi; 1 individual male extant, so ID in doubt); Conservation Assessment = E; US Status = E Melicope anisata (H. Mann) T. G. Hartley & B. C. Stone end; K

Melicope balloui (Rock) T. G. Hartley & B. C. Stone end; EM (between Olinda and ‘Ukulele) (extinct); Conservation Assessment = EX; US Status = E Melicope barbigera A. Gray end; K (Kōke‘e) Melicope christophersenii (H. St. John) T. G. Hartley & B. C. Stone end; O (Wa: Pu‘ukaua to Mt Ka‘ala); Conservation Assessment = R; US Status = E Melicope cinerea A. Gray end; O (Wa; Ko: Mānoa and Pālolo valleys)/ M (single collection); Conservation Assessment = R Melicope clusiifolia (A. Gray) T. G. Hartley & B. C. Stone end; K/ O/ Mo/ L/ M/ H Melicope cruciata (A. Heller) T. G. Hartley & B. C. Stone end; K (Kahōluamanu; Mt Wai‘ale‘ale); Conservation Assessment = R Melicope degeneri (B. C. Stone) T. G. Hartley & B. C. Stone end; K (Kōke‘e Stream; Hanakoa); Conservation Assessment = E; US Status = E Melicope elliptica A. Gray end; O (Wa)/ Mo (Kua Gulch)/ WM (Olowalu) Melicope feddei (H. Lév.) T. G. Hartley & B. C. Stone end; K Melicope haleakalae (B. C. Stone) T. G. Hartley & B. C. Stone end; EM (Olinda Flume Trail; ‘Ukulele); Conservation Assessment = R Melicope haupuensis (H. St. John) T. G. Hartley & B. C. Stone end; K (Hā‘upu); Conservation Assessment = E; US Status = E Melicope hawaiensis (Wawra) T. G. Hartley & B. C. Stone end; Mo/ L/ M/ H; Conservation Assessment = V Melicope hiiakae (B. C. Stone) T. G. Hartley & B. C. Stone end; O (Ko); Conservation Assessment = V; US Status = E Melicope hosakae (H. St. John) W. L. Wagner & R. K. Shannon end; O (Ko: Wiliwilinui to Pūpūkea) Melicope kaalaensis (H. St. John) T. G. Hartley & B. C. Stone end; O (Wa: Pu‘ukaua to Piko Trail) Melicope kavaiensis (H. Mann) T. G. Hartley & B. C. Stone end; K (Alaka‘i Swamp to Mt Wai‘ale‘ale) Melicope knudsenii (Hillebr.) T. G. Hartley & B. C. Stone end; K (Olokele Valley and Waimea)/ EM (Auwahi); Conservation Assessment = E; US Status = E Melicope lydgatei (Hillebr.) T. G. Hartley & B. C. Stone end; O (Ko: Hau‘ula-Kahana; Mānana-Waimano; PāloloWailupe); Conservation Assessment = E; US Status = E Melicope macropus (Hillebr.) T. G. Hartley & B. C. Stone end; K (Robinson’s summer house; Kahōluamanu) (extinct?); Conservation Assessment = EX? Melicope makahae (B. C. Stone) T. G. Hartley & B. C. Stone end; O (Wa: Pālāwai to Makaha Valley, ridges); Conservation Assessment = E; US Status = E Melicope molokaiensis (Hillebr.) T. G. Hartley & B. C. Stone end; Mo (Hanalilolilo to Oloku‘i)/ southeast WM (Manawainui to Pu‘ukukui)/ northeast EM Melicope mucronulata (H. St. John) T. G. Hartley & B. C. Stone end; Mo (Kūpā‘ia Gulch)/ EM (Pakiloi, south slope of Haleakalā, extinct); Conservation Assessment = E; US Status = E Melicope munroi (H. St. John) T. G. Hartley & B. C. Stone

APPENDIX

end; Mo (Kamalō)/ L (Lāna‘ihale); Conservation Assessment = E; US Status = E Melicope nealae (B. C. Stone) T. G. Hartley & B. C. Stone end; K (Kumuwela and Kahōluamanu) (extinct?); Conservation Assessment = EX? Melicope oahuensis (H. Lév.) T. G. Hartley & B. C. Stone end; O Melicope obovata (H. St. John) T. G. Hartley & B. C. Stone end; M (or possibly L) (extinct); Conservation Assessment = EX Melicope orbicularis (Hillebr.) T. G. Hartley & B. C. Stone end; WM (Pōhākea north to Honokōhua)/ EM (Makawao to Ko‘olau Gap) Melicope ovalis (H. St. John) T. G. Hartley & B. C. Stone end; EM (above Hāna; Kīpahulu Valley); Conservation Assessment = E; US Status = E Melicope ovata (H. St. John & E. P. Hume) T. G. Hartley & B. C. Stone end; northwest K/ O (Wa: Napepeiaulelo Gulch, N Pālāwai Gulch) Melicope pallida (Hillebr.) T. G. Hartley & B. C. Stone end; K (Hanapēpē; Koai‘e Canyon to Honopū and Limahuli valleys)/ O (S-cent. Wa: below Palikea; ‘Ēkahanui Gulch) (extinct); Conservation Assessment = E; US Status = E Melicope paniculata (H. St. John) T. G. Hartley & B. C. Stone end; K (upper Līhu‘e Ditch Trail; Wahiawa Bog; Lumaha‘i; Wailua River); Conservation Assessment = V; US Status = E Melicope peduncularis (H. Lév.) T. G. Hartley & B. C. Stone end; K (Kōke‘e)/ O/ east Mo (Ka‘aloa Peak)/ M Melicope pseudoanisata (Rock) T. G. Hartley & B. C. Stone end; EM (Kūhiwa Trail)/ H Melicope puberula (H. St. John) T. G. Hartley & B. C. Stone end; K (Kalalau Lookout to Pu‘uokila Lookout); Conservation Assessment = R; US Status = E Melicope quadrangularis (H. St. John & E. P. Hume) T. G. Hartley & B. C. Stone end; K (Wahiawa Bog); Conservation Assessment = EX?; US Status = E Melicope radiata (H. St. John) T. G. Hartley & B. C. Stone end; H (Kohala Mts to Puna District and SW to Hawaiian Ocean View Estates) Melicope reflexa (H. St. John) T. G. Hartley & B. C. Stone end; east Mo; Conservation Assessment = E; US Status = E Melicope rotundifolia (A. Gray) T. G. Hartley & B. C. Stone end; O (Ko: ‘Ōpae‘ula to Niu Valley) Melicope saint-johnii (E. P. Hume) T. G. Hartley & B. C. Stone end; O; Conservation Assessment = E; US Status = E Melicope sandwicensis (Hook. & Arn.) T. G. Hartley & B. C. Stone end; O (Wa; Ko: Kahana Iki to Waimalu); Conservation Assessment = R Melicope sessilis (H. Lév.) T. G. Hartley & B. C. Stone end; Mo (Waikolu to Kaunuohua)/ WM (Honokōwai to Pu‘ukukui)/ EM (Olinda to ‘Ōpana Gulch) Melicope volcanica (A. Gray) T. G. Hartley & B. C. Stone end; Mo (Wailau Valley)/ L/ EM/ H Melicope waialealae (Wawra) T. G. Hartley & B. C. Stone end; K (Wahiawa Bog; Alaka‘i Swamp to Mt Wai‘ale‘ale) Melicope wailauensis (H. St. John) T. G. Hartley & B. C. Stone end; Mo (Kukuinui Ridge in Wailau Valley) (extinct?); Conservation Assessment = EX?

Melicope wawraeana (Rock) T. G. Hartley & B. C. Stone end; K (Wahiawa Bog)/ O Melicope zahlbruckneri (Rock) T. G. Hartley & B. C. Stone end; H (Kīpukapuaulu; Moa‘ula, near Glenwood); Conservation Assessment = E; US Status = E Platydesma cornuta Hillebr. var. cornuta end; O (Ko); Conservation Assessment = R; US Status = E var. decurrens B. C. Stone end; O (Wa); Conservation Assessment = R; US Status = E Platydesma remyi (Sherff) O. Deg., I. Deg., Sherff & B. C. Stone end; H (Hāmākua-Kohala); Conservation Assessment = V; US Status = PE Platydesma rostrata Hillebr. end; K; Conservation Assessment = R; US Status = E Platydesma spathulata (A. Gray) B. C. Stone end; K/ O/ M/ H Zanthoxylum dipetalum H. Mann var. dipetalum end; K/ O/ Mo (Kamiloloa)/ H; Conservation Assessment = R var. tomentosum Rock end; H (Pu‘uwa‘awa‘a); Conservation Assessment = E; US Status = E Zanthoxylum hawaiiense Hillebr. end; K/ Mo/ L (extinct)/ M/ H; Conservation Assessment = E; US Status = E Zanthoxylum kauaense A. Gray end; K/ O/ Mo/ L/ M/ H Zanthoxylum oahuense Hillebr. end; O (Ko); Conservation Assessment = R; US Status = E Salicaceae Xylosma crenatum (H. St. John) H. St. John end; K (Mōhihi Stream; Nu‘alolo Trail); Conservation Assessment = E; US Status = E Xylosma hawaiiense Seem. end; K/ O/ Mo/ L/ M/ H Santalaceae Exocarpos gaudichaudii A. DC. end; Ni/ O/ Mo/ L/ M/ Ka/ H; Conservation Assessment = R Exocarpos luteolus C. N. Forbes end; K; Conservation Assessment = E; US Status = E Exocarpos menziesii Stauffer end; L (Kaiholena Gulch)/ leeward H (Kahuku Ranch and Mauna Loa Strip north to Hualālai and Pu‘ukapele) Korthalsella complanata (Tiegh.) Engl. ind; K/ O/ Mo/ L/ M/ H Korthalsella cylindrica (Tiegh.) Engl. end; O/ Mo/ L/ M/ H Korthalsella degeneri Danser end; O (Wa: Piko Trail; Mākua Valley); Conservation Assessment = R; US Status = E Korthalsella latissima (Tiegh.) Danser end; K/ O (Wa: Mt Ka‘ala)/ H (Waiākea Forest Reserve, Mauna Loa)

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Korthalsella platycaula (Tiegh.) Engl. ind; K/ O/ L (Kapu‘a Gulch)/ EM (Waikamoi Trail) Korthalsella remyana Tiegh. end; K/ O/ Mo/ L/ M/ H Santalum ellipticum Gaudich. end; La (extinct)/ HI (K [r]; Ka [ex]; H [Kohala Mts; Pu‘uwa‘awa‘a; Pu‘upapapa]) Santalum freycinetianum Gaudich. end; O Santalum haleakalae Hillebr. var. haleakalae end; EM (Haleakalā); Conservation Assessment = R var. lanaiense (Rock) Harbaugh end; Mo/L/ M; Conservation Assessment = E; US Status = E Santalum involutum H. St. John end; K; Conservation Assessment = E Santalum paniculatum Hook. & Arn. var. paniculatum end; H var. pilgeri (Rock) Stemmerm. end; H Santalum pyrularium A. Gray end; K Sapindaceae Alectryon macrococcus Radlk. var. auwahiensis G. Linney end; EM; Conservation Assessment = E; US Status = E var. macrococcus end; K/ O (Wa; Ko: Kīpapa Gulch)/ Mo/ M (Honokōwai Ditch Trail and Kahakapao Gulch); Conservation Assessment = E; US Status = E Dodonaea viscosa Jacq. ind; HI Sapindus oahuensis Hillebr. ex Radlk. end; northwest K/ O (Wa; Ko: Waimalu to Niu valleys) Sapindus saponaria L. ind; H (Hualālai; Mauna Loa; Kīlauea) Sapotaceae Planchonella sandwicensis (A. Gray) Pierre end; K/ O/ Mo/ L/ M/ H Sideroxylon polynesicum (Hillebr.) Smedmark & Anderb. ind; K/ O/ Mo/ L/ M; Conservation Assessment = VH Scrophulariaceae Myoporum sandwicense A. Gray ind; Ni/ K/ O/ Mo/ L/ M/ H Myoporum stellatum (G. L. Webster) O. Deg. & I. Deg. end; O (Barbers Point); Conservation Assessment = E Solanaceae Lycium sandwicense A. Gray ind; HI Nothocestrum breviflorum A. Gray end; H (Ka‘ū District north to Waimea); Conservation Assessment = E; US Status = E

Nothocestrum latifolium A. Gray end; K/ O/ Mo/ L/ M; Conservation Assessment = R; US Status = C Nothocestrum longifolium A. Gray end; K/ O/ Mo/ L/ M/ H Nothocestrum peltatum Skottsb. end; K (Kalalau Lookout; Kumuwela; Nu‘alolo; Mākaha Valley; west side of Waimea drainage); Conservation Assessment = E; US Status = E Solanum americanum Mill. ind; Ku/ Mi/ PH/ Li/ La/ N/ HI ind? Solanum incompletum Dunal end; K/ Mo/ L/ M/ H (extant only on H); Conservation Assessment = E; US Status = E Solanum nelsonii Dunal end; Ku/ Mi (extinct)/ PH/ La (extinct)/ N/ Ni/ K (extinct)/ O (extinct)/ Mo (Mo‘omomi to ‘Īlio Pt)/ M/ H; Conservation Assessment = V; US Status = C Solanum sandwicense Hook. & Arn. end; K/ O (ex); Conservation Assessment = E; US Status = E Thymelaeaceae Wikstroemia bicornuta Hillebr. end; L/ WM (Mt ‘Eke); Conservation Assessment = R Wikstroemia forbesii Skottsb. end; east Mo Wikstroemia furcata (Hillebr.) Rock end; K Wikstroemia hanalei Wawra end; north K (Wai‘oli, Moloa‘a, and Ho‘olulu valleys) (extinct?); Conservation Assessment = EX? Wikstroemia monticola Skottsb. end; EM Wikstroemia oahuensis (A. Gray) Rock var. oahuensis end; K/ O/ Mo/ L/ M var. palustris (Hochr.) Peterson end; K Wikstroemia phillyreifolia A. Gray end; H Wikstroemia pulcherrima Skottsb. end; H (South Kohala and North Kona districts) Wikstroemia sandwicensis Meisn. end; H Wikstroemia skottsbergiana Sparre end; K (Wahiawa Mts; Hanalei and Kauhao valleys); Conservation Assessment = R Wikstroemia uva-ursi A. Gray var. kauaiensis Skottsb. end; K var. uva-ursi end; O/ Mo/ M Wikstroemia villosa Hillebr. end; WM (extinct?), EM (Ko‘olau FR); Conservation Assessment = E; US Status = PE Urticaceae Boehmeria grandis (Hook. & Arn.) A. Heller end; K/ O/ Mo/ L/ M/ H

APPENDIX

Hesperocnide sandwicensis (Wedd.) Wedd. end; H (plateau between Hualālai, Mauna Loa, and Mauna Kea) Neraudia angulata R. S. Cowan var. angulata end; O (Wa); Conservation Assessment = E; US Status = E var. dentata O. Deg. & R. S. Cowan end; O (Wa); Conservation Assessment = E; US Status = E Neraudia kauaiensis (Hillebr.) R. S. Cowan end; K; Conservation Assessment = R Neraudia melastomifolia Gaudich. end; K/ O/ Mo (Wāwā‘ia Gulch)/ WM (Olowalu; ‘Īao Valley) Neraudia ovata Gaudich. end; leeward H; Conservation Assessment = E; US Status = E Neraudia sericea Gaudich. end; Mo/ L/ M/ Ka (extinct); Conservation Assessment = E; US Status = E Pilea peploides (Gaudich.) Hook. & Arn. ind; HI (Ni: Lehua Islet) Pipturus albidus (Hook. & Arn.) A. Gray end; K/ O/ Mo/ L/ M/ H Pipturus forbesii Krajina end; EM (Kaupō Gap east to Ko‘olau Gap) Pipturus kauaiensis A. Heller end; K Pipturus ruber A. Heller end; K Touchardia latifolia Gaudich. end; K/ O/ Mo/ L/ M/ H Urera glabra (Hook. & Arn.) Wedd. end; K/ O/ Mo/ L/ M/ H Urera kaalae Wawra end; O (Wa: Napepeiauolelo Gulch; ‘Ēkahanui near Pu‘ukaua; Kalua‘a near Pu‘uhāpapa); Conservation Assessment = E; US Status = E Violaceae Isodendrion hosakae H. St. John end; H (Nohonaohae and Pu‘upapapa; South Kohala District); Conservation Assessment = E; US Status = E Isodendrion laurifolium A. Gray end; K (Miloli‘i to Ku‘ia valleys; Pāpa‘alāi; Hā‘upu)/ O (Wa: Makaleha; Mākaha; Ko: Wailupe and Mānoa valleys); Conservation Assessment = E; US Status = E Isodendrion longifolium A. Gray end; K (Hanapēpē; Wahiawa Mts; Hā‘upu; Limahuli; Wainiha; and Hanakāpī‘ai)/ O (Wa: Palikea; Pu‘upane); Conservation Assessment = V; US Status = T Isodendrion pyrifolium A. Gray end; Ni/ O (Wa: Mt Ka‘ala)/ Mo/ L/ M/ H (Hualālai) extant only on H; Conservation Assessment = E; US Status = E Viola chamissoniana Ging. subsp. chamissoniana end; O (Wa); Conservation Assessment = E; US Status = E subsp. robusta (Hillebr.) W. L. Wagner, D. R. Herbst & Sohmer end; Mo

subsp. tracheliifolia (Ging.) W. L. Wagner, D. R. Herbst & Sohmer end; K/ O/ Mo/ M Viola helenae C. N. Forbes & Lydgate end; K (Wahiawa Stream); Conservation Assessment = E; US Status = E Viola kauaensis A. Gray var. kauaensis end; K (Waineki to Alaka‘i Swamp)/ O (Ko: between Kawainui and Kaipapa‘u) var. wahiawaensis C. N. Forbes end; K (Wahiawa Bog; Mt Wai‘ale‘ale); Conservation Assessment = E; US Status = E Viola lanaiensis W. Becker end; L (Lāna‘ihale)/ WM (Helu Peak); Conservation Assessment = E; US Status = E Viola maviensis H. Mann end; Mo (Pēpē‘ōpae Bog)/ M/ H (Kohala Mts) Viola oahuensis C. N. Forbes end; O (Ko: Kawailoa to Pālolo); Conservation Assessment = E; US Status = E Viola wailenalenae (Rock) Skottsb. end; K (Alaka‘i Swamp) Zygophyllaceae Tribulus cistoides L. ind; Ku/ Mi/ PH/ Li/ La/ FF/ N/ HI MONOCOTS

Araceae Spirodela polyrhiza (L.) Schleid. ind; K/ O (Ko)/ M/ H Arecaceae Pritchardia arecina Becc. end; EM (north slope Haleakalā) Pritchardia beccariana Rock end; H (Mauna Loa) Pritchardia flynnii Lorence & Gemmill end; K (Wahiawa Drainage, N Fork of Wailua River); Conservation Assessment = T Pritchardia forbesiana Rock end; WM (Honokōhau drainage; Mt ‘Eke)/ easternmost M; Conservation Assessment = R Pritchardia glabrata Becc. & Rock end; L/ WM (‘Īao Valley); Conservation Assessment = V Pritchardia gordonii Hodel end; H (Kohala Mts); Conservation Assessment = E Pritchardia hardyi Rock end; K (Powerline Trail); Conservation Assessment = R; US Status = E Pritchardia hillebrandii Becc. end; rocky islets of Huelo and Mōkapu off north coast of Moloka‘i Pritchardia kaalae Rock end; O (Wa: Mt Ka‘ala; Mākua-Mākaha Ridge);

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Conservation Assessment = E; US Status = E Pritchardia lanigera Becc. end; H (Kohala Mts; Waimea; S slope Mauna Loa; Nā‘ālehu; Mauna Kea between Honoka‘a and Pā‘auhau); Conservation Assessment = R; US Status = PE Pritchardia lowreyana Rock end; eastern Mo Pritchardia maideniana Becc. end; leeward H; Conservation Assessment = E; US Status = E Pritchardia martii (Gaudich.) H. Wendl. end; O (Ko) Pritchardia minor Becc. end; K (western Nāpali Coast, Kōke‘e, and Waimea Canyon) Pritchardia munroi Rock end; leeward Mo; WM; Conservation Assessment = E; US Status = E Pritchardia napaliensis H. St. John end; K (Nāpali Coast); Conservation Assessment = E; US Status = E Pritchardia perlmanii Gemmill end; K (Makaleha Mts; Power Line Trail; Wainiha Valley; Wai‘oli Valley); Conservation Assessment = E Pritchardia remota Becc. end; N (East and West Palm valleys)/ Ni (Ka‘ali Cliff); Conservation Assessment = E; US Status = E Pritchardia schattaueri Hodel end; H (Ho‘omau Ranchlands of Pāpā); Conservation Assessment = E; US Status = E Pritchardia viscosa Rock end; K (Powerline Trail); Conservation Assessment = E; US Status = E Pritchardia waialealeana Read end; K (Powerline Trail) Pritchardia woodii Hodel end; EM (Waiho‘i and Kīpahulu valleys); Conservation Assessment = E Asparagaceae Pleomele aurea (H. Mann) N. E. Br. end; K Pleomele auwahiensis H. St. John end; central Mo/ leeward M Pleomele fernaldii H. St. John end; L; Conservation Assessment = E; US Status = PE Pleomele forbesii O. Deg. end; O (Wa; Ko: Kawela; Wilhelmina Rise); Conservation Assessment = R; US Status = E Pleomele halapepe H. St. John end; O Pleomele hawaiiensis O. Deg. & I. Deg. end; leeward H; Conservation Assessment = E; US Status = E Asteliaceae Astelia argyrocoma A. Heller ex Skottsb. end; K Astelia menziesiana Sm. end; K/ O/ Mo/ L/ M/ H

Astelia waialealae Wawra end; K (Mt Wai‘ale‘ale; Alaka‘i Swamp); Conservation Assessment = E; US Status = E Cyperaceae Bolboschoenus maritimus (L.) Palla ind; Ni/ K/ O/ Mo/ M/ H Carex alligata Boott end; K/ O/ Mo/ M/ H Carex echinata Murray ind; K/ M/ H Carex kauaiensis R. W. Krauss end; K Carex macloviana d’Urv. subsp. subfusca (Boott) T. Koyama ind; M/ H Carex meyenii Nees ind; HI (Ka [extinct]) Carex montis-eeka Hillebr. end; K/ Mo/ M Carex nealae R. W. Krauss end; M/ H ind? Carex wahuensis C. A. Mey. subsp. herbstii T. Koyama end; O (Ko: Moanalua Valley; single collection) (extinct?); Conservation Assessment = EX? subsp. rubiginosa (R. W. Krauss) T. Koyama end; K/ O/ Mo (Kamakou Preserve)/ L/ M/ H subsp. wahuensis end; K/ O/ Mo/ L/ M/ H Cladium jamaicense Crantz ind; K/ O/ Mo/ M/ H Cyperus cyperinus (Retz.) Suringar ind; K/ O/ Mo/ EM Cyperus fauriei Kük. end; Mo/ L (extinct)/ H; Conservation Assessment = R; US Status = E Cyperus hillebrandii Boeck. var. decipiens (Hillebr.) Kük. end; K/ O (Wa)/ L/ WM var. hillebrandii end; O/ L/ EM/ H Cyperus hypochlorus Hillebr. var. brevior Kük. end; K/ O/ Mo/ M var. hypochlorus end; K/ O/ Mo/ M/ H Cyperus javanicus Houtt. ind; Mi/ Ni/ K/ O/ Mo/ L/ M/ H Cyperus laevigatus L. ind; La/ Ni/ O/ Mo/ M/ H Cyperus neokunthianus Kük. end; WM (extinct); Conservation Assessment = EX Cyperus odoratus L. ind; K (extinct)/ O/ Mo (extinct)/ M (extinct)/ H (extinct); Conservation Assessment = R Cyperus pennatiformis Kük. var. bryanii Kük.

APPENDIX

end; northeast La; Conservation Assessment = E; US Status = E var. pennatiformis end; K (extinct?)/ O (extinct)/ M (extinct?)/ H (extinct?); Conservation Assessment = EX?; US Status = E Cyperus phleoides (Nees ex Kunth) Hillebr. var. hawaiiensis (H. Mann) Kük. end; Ni/ K/ O/ Mo/ L/ M var. phleoides end; K/ Mo/ L/ M/ Ka/ H Cyperus polystachyos Rottb. ind; Mi/ Ni/ K/ O/ Mo/ L/ M/ H Cyperus rockii Kük. end; K (Wai‘alae Valley) (extinct?); Conservation Assessment = EX? Cyperus sandwicensis Kük. end; K/ O/ Mo/ M Cyperus trachysanthos Hook. & Arn. end; Ni (extinct)/ K/ O/ Mo (extinct)/ L (extinct); Conservation Assessment = E; US Status = E Eleocharis calva Torr. ind; Ni/ O/ Ka ind? Eleocharis obtusa (Willd.) Schult. ind; K/ O/ Mo/ M/ H Fimbristylis cymosa R. Br. subsp. spathacea (Roth) T. Koyama ind; Mi/ La/ FF/ Ni/ K/ O/ WM/ H subsp. umbellato-capitata (Hillebr.) T. Koyama ind; Ku/ Mi/ La/ Ni/ K/ O/ Mo/ L/ M/ H Fimbristylis dichotoma (L.) Vahl ind; K/ O/ Mo/ L/ M/ H Fimbristylis hawaiiensis Hillebr. end; H (Ka‘ū Desert; Chain of Craters Rd, Hawai‘i Volcanoes NP); Conservation Assessment = R Gahnia aspera Spreng. subsp. globosa (H. Mann) J. Kern end; K / O Gahnia beecheyi H. Mann end; K/ O/ L/ Mo/ M/ H Gahnia vitiensis Rendle subsp. kauaiensis (Benl) T. Koyama end; K (Alaka‘i Swamp to Mt Wai‘ale‘ale) Machaerina angustifolia (Gaudich.) T. Koyama ind; K/ O/ Mo/ L/ M/ H Machaerina mariscoides (Gaudich.) J. Kern subsp. meyenii (Kunth) T. Koyama end; K/ O/ Mo/ L/ M/ H Morelotia gahniiformis Gaudich. end; K (Wahiawa)/ Mo/ L/ M/ H Oreobolus furcatus H. Mann end; K/ O/ Mo/ M Rhynchospora chinensis Nees & Meyen subsp. spiciformis (Hillebr.) T. Koyama ind; K/ O (Ko‘olau Mts)/ Mo/ M/ H Rhynchospora rugosa (Vahl) Gale subsp. lavarum (Gaudich.) T. Koyama ind; K/ O/ Mo/ M/ H Rhynchospora sclerioides Hook. & Arn. ind; K/ O/ Mo/ L/ M/ H

Schoenoplectiella juncoides (Roxb.) Lye ind; K/ H (Kohala Mts) Schoenoplectus tabernaemontani (C. C. Gmel.) Palla ind; Ni/ K/ O/ Mo/ H Scleria testacea Nees ind; Mo/ WM (Pu‘ukukui)/ H (Hilo; Kīlauea) Uncinia brevicaulis Thouars ind; EM (Ke‘anae to Pōhakupālaha) Uncinia uncinata (L. f.) Kük. ind; K/ Mo/ M/ H Hydrocharitaceae Halophila decipiens Ostenf. ind; Mi (inner harbor, between Cargo Pier and Turtle Beach)/ O (along Kahala Beach coast)/ H (off Kohala Coast) Halophila hawaiiana Doty & B. C. Stone end; Mi/ PH/ K/ O/ Mo/ M Iridaceae Sisyrinchium acre H. Mann end; EM/ H; Conservation Assessment = R Joinvilleaceae Joinvillea ascendens Gaudich. ex Brongn. & Gris subsp. ascendens end; K/ O/ Mo/ M/ H; Conservation Assessment = R; US Status = C Juncaceae Luzula hawaiiensis Buchenau var. glabrata (Hillebr.) O. Deg. & I. Deg. end; K/ Mo/ M var. hawaiiensis end; K/ L/ M/ H var. oahuensis (O. Deg. & Fosberg) O. Deg. & I. Deg. end; O (Wa) Orchidaceae Anoectochilus sandvicensis Lindl. end; K/ O/ Mo/ L/ M/ H; Conservation Assessment = R Liparis hawaiensis H. Mann end; K/ O/ Mo/ L/ M/ H; Conservation Assessment = R Platanthera holochila (Hillebr.) Kraenzl. end; K/ O (Ko) (extinct)/ Mo/ M; Conservation Assessment = E; US Status = E Pandanaceae Freycinetia arborea Gaudich. ind; K/ O/ Mo/ L/ M/ H Pandanus tectorius Parkinson ind; Ni/ K/ O/ Mo/ L/ M/ H ind? Poaceae Agrostis sandwicensis Hillebr. end; K/ O (reported by Hillebrand, 1888)/ M/ H

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APPENDIX

Calamagrostis expansa (Munro ex Hillebr.) Hitchc. end; M/ H (Kohala Mts); Conservation Assessment = R; US Status = C Calamagrostis hillebrandii (Munro ex Hillebr.) Hitchc. end; central Mo (reported by Hitchcock, 1922; extinct)/ WM (Pu‘ukukui); Conservation Assessment = V; US Status = PE Cenchrus agrimonioides Trin. var. agrimonioides end; O/ Mo/ L/ M/ H (extinct); Conservation Assessment = E; US Status = E var. laysanensis F. Br. end; Ku/ Mi/ La (extinct); Conservation Assessment = EX; US Status = E Chrysopogon aciculatus (Retz.) Trin. ind; K/ O/ Mo/ L/ WM/ H ind? Deschampsia nubigena Hillebr. end; K/ Mo/ M/ H Dichanthelium cynodon (Reichardt) C. A. Clark & Gould end; K/ O / Mo/ M Dichanthelium hillebrandianum (Hitchc.) C. A. Clark & Gould end; K/ O/ Mo/ M/ H Dichanthelium isachnoides (Munro ex Hillebr.) C. A. Clark & Gould end; K/ Mo/ M Dichanthelium koolauense (H. St. John & Hosaka) C. A. Clark & Gould end; O (Ko: Kaipapa‘u to Moanalua)/ WM (Mt ‘Eke); Conservation Assessment = R Digitaria setigera Roth ind; Ni/ K/ O/ Mo/ L/ M/ H Dissochondrus biflorus (Hillebr.) Kuntze ex Hack. end; K (extinct)/ O/ Mo/ L/ M/ H; Conservation Assessment = R Eragrostis atropioides Hillebr. end; O/ Mo/ L/ M/ H Eragrostis deflexa Hitchc. end; Mo (Kamakou Preserve)/ L (extinct)/ WM (Līhau)/ H; Conservation Assessment = R Eragrostis fosbergii Whitney end; O (Wa: Mt Ka‘ala to Kolekole Pass); Conservation Assessment = E; US Status = E Eragrostis grandis Hillebr. end; HI exc. Ni Eragrostis leptophylla Hitchc. end; L/ M/ Ka/ H Eragrostis mauiensis Hitchc. end; L/ M (Wailuku) (extinct); Conservation Assessment = EX Eragrostis monticola (Gaudich.) Hillebr. end; Mo/ L/ M/ H Eragrostis paupera Jedwabn. ind; Ku/ Mi/ PH/ FF/ O (Barbers Pt; extinct) Eragrostis variabilis (Gaudich.) Steud. end; Ku/ Mi/ PH/ Li/ La/ N/ HI Festuca aloha Catalán, Soreng & P. M. Peterson end; K; Conservation Assessment = V Festuca hawaiiensis Hitchc. end; EM (‘Ulupalakua, single collection, ex)/ H

(Pu‘uhuluhulu and Hualālai); Conservation Assessment = R; US Status = C Festuca molokaiensis Soreng, P. M. Peterson & Catalán end; Mo (Kūpā‘ia Gulch) ; US Status = PE Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult. ind; HI ind? Isachne distichophylla Munro ex Hillebr. end; K/ O/ Mo/ L/ M/ H Isachne pallens Hillebr. end; K/ O Ischaemum byrone (Trin.) Hitchc. end; K/ O/ Mo/ M/ H; Conservation Assessment = E; US Status = E Lachnagrostis filiformis (G. Forst.) Trin. ind; K/ O/ Mo/ L/ M /Ka/ H Lepturus repens (G. Forst.) R. Br. ind; Ku/ Mi/ PH/ Li/ La/ FF/ Ni (Lehua Islet) Panicum beecheyi Hook. & Arn. end; K/ O/ M Panicum fauriei Hitchc. var. carteri (Hosaka) Davidse end; O (Mokoli‘i Islet)/ Mo/ L/ M; Conservation Assessment = E; US Status = E var. fauriei end; O/ Mo/ L/ M/ Ka/ H var. latius (H. St. John) Davidse end; HI (Ni: Lehua Islet) Panicum konaense Whitney & Hosaka end; K (Waimea District)/ Mo/ M/ H Panicum lineale H. St. John end; K (Wainiha to Kalalau valleys); Conservation Assessment = R Panicum longivaginatum H. St. John end; H (South Hilo District, Power Line Rd.); Conservation Assessment = R Panicum nephelophilum Gaudich. end; K/ O/ Mo/ L/ M/ H Panicum niihauense H. St. John end; Ni (extinct)/ K (Polihale); Conservation Assessment = E; US Status = E Panicum pellitum Trin. end; Ni/ K/ L/ M/ H Panicum ramosius Hitchc. end; Mo/ L/ Ka; Conservation Assessment = R Panicum tenuifolium Hook. & Arn. end; K/ O/ Mo/ L/ M/ H Panicum torridum Gaudich. end; Ne/ N/ HI Panicum xerophilum (Hillebr.) Hitchc. end; Ni/ O/ Mo/ L/ M/ Ka/ H Paspalum scrobiculatum L. ind; K/ O/ Mo/ M/ H ind? Poa mannii Munro ex Hillebr. end; K (Hanalei and Waimea districts; Olokele Gulch); Conservation Assessment = E; US Status = E Poa sandvicensis (Reichardt) Hitchc. end; K; Conservation Assessment = E; US Status = E Poa siphonoglossa Hack. end; K (Kōke‘e); Conservation Assessment = E; US Status = E

APPENDIX

Sporobolus virginicus (L.) Kunth ind; Mi/ La/ HI Trisetum glomeratum (Kunth) Trin. end; M/ H Trisetum inaequale Whitney end; L/ M Potamogetonaceae Potamogeton foliosus Raf. ind; K/ O/ M/ H ind? Potamogeton nodosus Poir. ind; O/ M ind?

Ruppiaceae Ruppia maritima L. ind; HI exc. Ka Smilacaceae Smilax melastomifolia Sm. end; K/ O/ Mo/ L/ M/ H Xanthorrhoeaceae Dianella sandwicensis Hook. & Arn. end; K/ O/ Mo/ L/ M/ H

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GLOSSARY

abaxial. Referring to the underside of a leaf or frond. acaulescent. Lacking a stem. achene. Dry indehiscent one-seeded fruit. achlorophyllous. Lacking chlorophyll. actinomorphic. Radially symmetrical, in contrast to zygomorphic. adaxial. Referring to the upper side of a leaf or frond. adherent. Joined with. adventitious roots. Roots arising from mature tissue where not normally expected such as from a stem. alkaloid. Naturally occurring nitrogenous compounds that are the active components of many drugs and pharmaceuticals, including nicotine, atropine, morphine, quinine, and theobromine. annual. Plant that germinates and completes its life cycle in a single year or growing season. anther. Pollen-bearing portion of a stamen containing 1 or 2 pollen sacs. anthocyanins. Water-soluble glucoside pigments ranging in color from purple and red to blue and violet. apical. At or near the apex. apomixis. Asexual seed production without fertilization. appressed. Pressed flat against another organ. arborescent. Treelike in growth habit. areole. Space enclosed by veins in a vein network. aril. Outgrowth of the stalk attaching an ovule to the ovary wall; often fleshy and colorful to attract animals, thus aiding in seed dispersal. awn. Slender bristle, usually terminal or dorsal in position, on the spikelet of many grasses. axil. Point between the stem and a leaf petiole. axillary. Located in or arising from an axil. berry. Fruit developed from a simple or compound ovary in which the ovary walls and inner tissues become enlarged and juicy, and containing 2 to many seeds. bilobed. Having 2 lobes. bipinnate. Twice pinnate with the lobes of the pinnae pinnate. bipinnatifid. Pinnatifid with lobes further divided. blade. Flat, expanded part of a plant structure such as a leaf or petal. bract. Specialized leaf associated with a flower or inflorescence. bulb. Short, vertical underground stem with modified leaves or thickened leaf bases that serves to store carbohydrates. bulblet. Small subsidiary bulb.

C 4 metabolism. Metabolic system in which there are two spatially separated carboxylation processes; this aids in water use efficiency and in allowing high rates of photosynthesis; common in lowland tropical grasses. caespitose. Growing in dense tufts; matted. calyx. Collective term for all the sepals of a flower. capitate. Headlike. capsule. Dry, dehiscent fruit consisting of 2 or more carpels. carpel. Ovule-bearing structure of a flower, consisting of the ovary, stigma, and style; several carpels may be fused to form a compound ovary or gynoecium. caryopsis. Dry fruit of grasses, derived from a single-celled superior ovary; the seed coat and fruit wall are fused; compare with achene, in which they remain separate. caudex (plural caudices). A thick trunklike rhizome, especially in reference to tree ferns. cauliferous. Producing flowers and fruits on the trunk or primary branches. cauline. Growing on the stem. chaff. Thin scalelike bracts, often common in the Asteraceae. chartaceous. Papery in texture. chasmogamous. Referring to open flowers allowing external pollination to take place, as contrasted with cleistogamous. circinnate vernation. Uncoiling of new leaves from the base toward the apex, as in ferns. cleistogamous. A flower that does not open and is thus obligatorily self-pollinated, as in the Violaceae. coherent. Sticking together, applied only to like parts. compound leaf. Leaf with 2 or more separate leaflets; these may be palmate or pinnately arranged. cordate. Heart shaped. coriacious. Leathery in texture. corm. Short, vertical underground stems that function to store carbohydrates and are enclosed by dry, scalelike leaves. corolla. Collective term for all the petals of a flower. corymb. A racemelike inflorescence in which the stalks of the individual flowers vary in length, resulting in a flat-topped or convex inflorescence. costa (plural costae). A prominent rib or vein such as the midrib of a fern pinna. crassulacean acid metabolism. Metabolic system used by many succulent plants and epiphytes in which there are two separate carboxylation processes, with one fixing carbon in the dark and storing this as organic acids and then

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GLOSSARY

breaking down these compounds during the day in the same tissues to refix the carbon for use in the light reactions of photosynthesis. creeping. Growing horizontally along the ground. crenulate. Having shallow rounded teeth along the leaf ’s margin. cruciform. Cross shaped. culm. The aerial stem of a grass or sedge. cyme. Determinant inflorescence with the terminal flower blooming first. deciduous. Tissues shed after the end of their normal function or life, as with leaves that fall at the beginning of the dormant season. decumbent. Growth habit with a portion of the stem trailing along the ground without rooting and with the upper part of the stem upright and erect. decussate. Arrangement of opposite leaves with each succeeding pair arrayed at right angles to the previous pair; compare with distichous. dehiscent. Opening at maturity. deltate. Triangular shaped. dichotomous. Regularly forking with branches at each fork equal in size. dimorphic. Having 2 distinct forms, such as having different juvenile and adult leaves or sterile and fertile flowers. dioecious. Having male and female flowers of the same species on different individual plants. diploid. Having cells with 2 homologous copies of each chromosome. disarticulate. To fall or be shed. discoid. Disklike, or having disk flowers, tubular perfect flowers as in the Asteraceae. disk. Fleshy outgrowth from the receptacle surrounding the base of the ovary. distal. Toward the tip, in contrast to basal. distichous. Arrangement of opposite leaves with each succeeding pair arrayed in the same plane as the previous pair; compare with decussate. domatium (plural domatia). A small pit or cavity, usually on a leaf, that forms a habitat for insects, mites, fungi, or bacteria; common in Psychotria. drupaceous. Resembling a drupe in general appearance but not necessarily with a drupe structure. drupe. Fleshy fruit derived from a single carpel and containing a hardened pit enclosing a seed, such as in the peach, olive, mango, or cherry. elliptical. Having the shape of an elongated circle. endosperm. The storage tissue of an angiosperm seed. entire. Having a smooth leaf margin without teeth. epiphyte. Plant growing attached to the trunk or branch of another plant or object above the ground but not deriving resources directly from the host plant. erect. Upright in growth. fertile. Capable of reproduction; in ferns and fern relatives, those fronds or branches that bear sporangia. filament. The stalk of a stamen. filiform. Slender, threadlike. floret. Small flower that is part of a dense flower cluster; in grasses, the spikelet. foliaceous. Leaflike.

frond. The photosynthetic organ of a fern, including both the blade and stipe. Also a palm leaf. fusiform. Spindle shaped; thickest in the middle and tapering to both ends. gametophyte. Sexual and inconspicuous haploid stage of the life cycle of higher plants. gemma (plural gemmae). Asexual reproductive organ that becomes detached from the parent to establish a new plant; termed proliferations in ferns. glabrous. Lacking hairs or glands. glandular. Having glands or functioning as a gland, as in glandular hairs. glaucous. Covered with a fine waxy coating. glumes. Pair of small bracts found at the base of the spikelet in grasses; also the bract subtending each flower in sedges. globose. Spherical in shape. gynodioecious. With only female flowers produced on one plant and perfect flowers on another. gynoecium. A collective term for all of the carpels of a flower. gynomonoecious. Having female flowers and perfect but no male flowers on the same plant. halophytic. Tolerating high levels of salinity. haustorium (plural haustoria). Specialized roots of parasitic plants able to penetrate the roots or tissues of a host plant and tap resources. head. Dense indeterminate inflorescence of sessile or subsessile flowers crowded closely together. heterophyllous. Having leaves of different sizes or shapes. hypanthium. Floral structure consisting of the bases of the sepals, petals, and stamens fused together; its presence is diagnostic of certain families, including the Rosaceae and Fabaceae. imbricate. Sepals, petals, or other organs arranged in an overlapping manner like shingles on a roof. indehiscent. Remaining closed at maturity. indusium (plural indusia). Thin epidermal outgrowth from a fern blade that covers the sorus. inferior ovary. Ovary with other flower parts attached to its apex. inflorescence. Flower cluster or the arrangement of the flowers. involute. Having margins rolled over toward their upper surface, in contrast to revolute. lanceolate. Lance shaped, with its widest point between the middle and base and tapering toward the tip. latex. A liquid that may be white, other colors, or colorless produced in the tissues of some plants; often it is toxic. leaflet. Smallest unit of a compound leaf. lemma. One of the pairs of bracts that subtends the spikelet in grass flowers; the lemma has its back at the outside of the spikelet, while the palea has its back to the axis of the spikelet. lenticel. Spongy area in the bark or outer stem that allows the interchange of gases between the atmosphere and the internal tissues. liana. Woody vine. ligule. Strap-shaped structure, such as the corolla of a ray floret of some species of Asteraceae; an outgrowth at the junction of the blade and sheath of the leaves of most grasses and a few sedges; the small flap of leaf tissue on the adaxial leaf surface just above the sporangia in Isoëtes and Selaginella.

GLOSSARY

lodicules. Tiny scale in grasses that may represent a vestigial perianth. marginal. Borne on the margin of an organ. membranous. Thin and flexible and relatively translucent. midrib. Central or primary rib or vein of a leaf. monoecious. Having separate male and female flowers on the same plant. mycorrhiza (plural mycorrhizae). Symbiotic fungus that lives associated with or within the tissues of a vascular plant root and aids in the uptake of nutrients and water. nectary (plural nectaries). Structure that produces nectar, usually on a flower but also present on some vegetative organs. neotropics. New world tropics. nerve. Prominent lateral vein of a leaf or other organ. nut. Hard, dry, indehiscent 1-seeded fruit with a hard shell. nutlet. Dry, indehiscent 1-seeded half carpel in the Boraginaceae and Lamiaceae. oblanceolate. Lance shaped, with the widest portion between the middle and the tip. obovate. Ovate, with the widest portion toward the tip. oligotrophic. Low in nutrients and nutrient availability. ovary. Structure that encloses the ovules of flowering plants; typically the expanded basal part of the pistil. ovate. Egg shaped. palea. One of the pairs of bracts that subtends the spikelet in grass flowers; the palea has its back to the axis of the spikelet, while the lemma has its back at the outside of the spikelet. paleotropics. Old world tropics. palmate. Having lobes, leaflets, or veins radiating from a common point, as with the fingers of a hand. panicle. A branched indeterminate inflorescence with pedicellate flowers borne on secondary branches. paniculate. Having an inflorescence in the form of a panicle. pantropical. Distributed in both the neotropics and paleotropics. papilla (plural papillae). A short nipplelike epidermal projection. papillate. Covered with papillae. pappus. Hairs, scales, and/or bristles that crown the ovary of flowers in the Asteraceae; these are a highly modified calyx. parasitic. Deriving nourishment in the form of carbohydrates, nutrients, and/or water from another plant to the detriment of that plant. pedicel. Stalk of a single flower in an inflorescence; the stalk of a spikelet in grasses. pedicellate. Having a pedicel. peduncle. Stalk of an inflorescence or a solitary flower. peltate. Round or oval in shape and attached by a central stalk. pendulous. Hanging loosely. perfect flower. Flower with both functioning male and female organs. perianth. Collective term for the combined sepals and petals of a flower. persistent. Remaining attached after the normal function or life of a tissue is completed. petal. Inner whorl of the perianth, often brightly colored to attract pollinators; the petals collectively are the corolla. petiole. Stalk of a leaf that connects the blade to the stem.

phyllode. Flattened petiole that takes on the function of a leaf as a photosynthetic organ. pinna (plural pinnae). Primary division of a compound fern blade. pinnate. Having 2 rows of leaflets arranged along a common axis. pinnatifid. Deeply lobed along an axis but not cut all of the way to the midrib. pinnatisect. Deeply cut along an axis all of the way to the midrib. pinnule. Secondary unit of division in a twice-divided fern blade. pistil. Female organ of a flower consisting of 1 or more carpels and typically differentiated into ovary, style, and stigma. pistillate flower. A flower lacking functional anthers and thus female. polygamodioecious. Functionally dioecious, with some of the flowers perfect on otherwise staminate or pistillate plants. prickle. Sharp outgrowth from the epidermis or bark. procumbent. Trailing along the ground but not rooting at the nodes. proliferation. Plantlet or bulblet growing on the frond or rhizome of a fern. prop root. Adventitious root that arises from the stem and angles down to the soil to help support the plant stem. prostrate. Growing flat along the ground but not rooting at the nodes. protuberance. Tissue that bulges out of or extends from the organ below. pseudobulb. Thickened stems of some orchids, used as a storage organ derived from the part of a stem between 2 leaf nodes. pseudodichotomous. Appearing to be dichotomous but not truly, even in branching. pubescent. Having hairs. punctate. Dotted with small pits that may be translucent or glandular. quadrangular. Square in cross section. raceme. Unbranched indeterminate inflorescence with stalked flowers arranged along the axis. racemose. Having an inflorescence with the form of a raceme. rachis (plural rachises). Axis or midrib of a leaflet or fern blade. radiate. Having ray flowers, as in the Asteraceae. ray floret. Strap-shaped flowers that comprise all of the marginal flowers in the head of Asteraceae; one of the branches of an umbel. receptacle. End of the peduncle or pedicel where the flower parts are attached; in ferns and fern relatives, the thickened tissue on which the sporangia are borne. recurved. Curved or curled downward or backward. reflexed. Abruptly bent or curved backward. resin duct. Tubelike intercellular structures or glands that, in certain plants, secrete resins, especially when wounded, resins that are a mixture of high molecular weight compounds such as terpenoids. reticulate. Forming a network, as with the veins on a leaf. retrorse. Pointing backward. revolute. Having lateral margins that are curled toward the lower surface; compare with involute. rhizomatous. Bearing rhizomes.

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GLOSSARY

rhizome. Underground horizontal stem of a plant, which, by breaking or branching, acts as a form of vegetative propagation. rosette. Circular arrangement of leaves radiating from a single point and reaching a common height or forming a spherical shape. rosulate. Arranged as in a basal rosette. rupestral. Growing on rocks. sagittate. Arrowhead shaped. saprophyte. Plants that live on dead organic matter; commonly used to include mycotropic plants that obtain their food through a symbiotic fungal associate. scale. Flat multicellular outgrowth from a plant tissue. scandent. Loosely climbing or leaning for support on other plants. schizocarp. Fruit that splits into 2 or more separate carpels at maturity; the units may be dehiscent or indehiscent and dry or fleshy. sepal. Outermost ring of sterile floral parts, typically green in color; the sepals collectively are called the calyx. serrate. Toothed along the margin. sessile. Lacking a petiole or stalk and attached directly at the base. silicle. Fruit similar to a silique but as wide or wider than long; characteristic of the Brassicaceae. silique. Elongate capsule in which the 2 valves are deciduous from the persistent seed-bearing partition of the fruit; characteristic of the Brassicaceae. sorus (plural sori). Cluster of sporangia. spadix. A fleshy spike of small crowded flowers, especially in the Araceae. spathe. A large and usually showy bract enclosing a spadix, especially in the Araceae. spathulate. Spoon shaped; that is, rounded at the tip and narrowed at the base. spike. Unbranched indeterminate inflorescence with sessile flowers arranged along the axis. spikelet. One of the ultimate flower clusters in grasses and sedges consisting of 1 to many flowers and their subtending bracts. spine. Sharp-pointed structure; it is a modified leaf or stipule. sporangium (plural sporangia). Spore-bearing organ of ferns and fern relatives. sporocarp. Stalked globular organ containing the sporangia of aquatic ferns. sporophyte. Dominant asexual and diploid form of growth in nonflowering plants that produces spores. stalk. Short supporting axis, as with a leaf, fruit, or pinna. staminate flower. Having stamens but no functional female parts. staminode (plural staminodea). Modified stamen that does not produce pollen; often showy and petal-like in appearance. stellate. Star shaped, usually in reference to hairs. stipe. Petiole of a fern frond; the stalk of any structure but usually used only when a more precise term is not available. stipule. Outgrowth on either side of the base of a leaf petiole; stipules appear as glands, scales, hairs, spines, or leaflike structures. stolon. Creeping stem along the surface of the ground and usually rooting at its nodes. stoloniferous. Possessing stolons.

strobilus (plural strobili). Spike of fertile leaves with sporangia at the base, as present in the Lycopodiaceae and Selaginellaceae. style. Slender stalk that connects the stigma to the ovary. submarginal. Borne close to but not on the margin of an organ. suborbicular. Semiround in shape. subsessile. Almost but not perfectly sessile. subshrub. Perennial plant, woody at the base but herbaceous above. subtend. To be placed directly below and close. succulent. Having fleshy tissue. suffrutescent. Semiwoody. superior ovary. Ovary attached to the summit of the receptacle and distinct from other flower parts that arise below. sympatric. Species or taxa distributed with an overlapping geographic range. sympetalous. Having the petals joined, at least at their base. synangium (plural synangia). Cluster of sporangia fused with development, as with Psilotum and certain Marattiaceae. taproot. Enlarged primary plant root that grows vertically downward; it forms the axis from which other roots spread laterally. tendril. Coiling or twisting organ of a vine used for attachment, usually representing a modified stem, leaf, or leaf part. tepal. Undifferentiated sepals and petals. terete. Round in cross section. terminal. Arising or positioned at the end of a branch or inflorescence. ternate. Arranged in threes. tetraploid. Having cells with 4 sets of homologous chromosomes. thalloid. Resembling the undifferentiated tissue of a lower plant. thorn. Stiff, woody, modified stem. tomentose. Covered with matted wooly hairs. tomentum. Matted wooly hairs. trichome. General term for a hair, scale, or bristle. trifoliate. Having 3 leaflets. tuber. Thickened portion, usually at the end of a rhizome, typically serving as a storage or asexual reproduction organ. tubercle. Swelling or projection, usually distinctive in color or texture from the organ from which it is derived. tuberous root. A thickened portion of a root that functions as a tuber but is not stem tissue. tussock. A dense tuft of grass or similar grasslike plant. umbel. Inflorescence consisting of a number of short flower stalks (pedicels) that are equal in length and spread from a common point; characteristic of the Apiaceae but not restricted to that family. umbellate. Having an inflorescence that forms an umbel. uniseriate. Arranged in a single row. valve. One of the portions of the ovary wall into which a capsule separates at maturity; the portion of the anther wall covering the pore; in ferns, the lobe of an indusium. vascular bundle. Unit strand of xylem and phloem in a vascular plant. vein. Strand of vascular tissue in a leaf or fern blade. verticillate. Arranged in whorls. viscid. Sticky or glutinous. zygomorphic. Bilaterally symmetrical, in contrast to actinomorphic.

FURTHER READING

Hotchkiss, S., P. M. Vitousek, O. A. Chadwick, and J. Price. 2000. Climate cycles, geomorphological change, and the interpretation of soil and ecosystem development. Ecosystems 3:522–533. Juvik, J. O., and D. Nullet. 1995. Relationships between rainfall, cloud-water interception, and canopy through fall in a Hawaiian montane forest. In L. S. Hamilton, J. O. Juvik, and F. N. Scantena (eds.), Tropical Montane Cloud Forests. Springer Verlag, Berlin, pp. 165–182. Juvik, J. O., D. Nullet, P. Banko, and K. Hughes. 1995. Forest climatology near the tree line in Hawaii. Agricultural and Forest Meteorology 66:159–172. Juvik, S. P., and J. O. Juvik (eds.). 1999. Atlas of Hawai‘i, 3rd ed. University of Hawai‘i Press, Honolulu. Kolivras, K. N., and A. C. Comrie. 2007. Regionalization and variability of precipitation in Hawaii. Physical Geography 28:76–96. Lintner, B. R., W. Buermann, C. D. Koven, and I. Y. Fung. 2006. Seasonal circulation and Mauna Loa CO2 variability. Journal of Geophysical Research: Atmospheres 111:doi:10.1111/j.1365–2486.2007.01420.x. MacDonald, G. A. 1983. Volcanoes in the Sea: The Geology of Hawaii, 2nd ed. University of Hawai‘i Press, Honolulu. Nullet, D., J. O. Juvik, and A. Wall. 1995. A Hawaiian mountain climate cross-section. Climate Research 5:131–137. Price, J. P., and D. A. Clague. 2002. How old is the Hawaiian biota? Geology and phylogeny suggest recent divergence. Proceedings of the Royal Society of London Series B: Biological Sciences 269:2429–2435. Price, J. P., and D. L. Elliott-Fisk. 2004. Topographic history of the Maui Nui complex, Hawai‘i, and its implications for biogeography. Pacific Science 58:27–54. Rooney, J. J., P. Wessel, R. Hoeke, J. Weiss, J. Baker, F. Parrish, C. H. Fletcher, J. Chojnacki, M. Garcia, R. Brainard, and Peter Vroom. 2008. Geology and geomorphology of coral reefs in the Northwestern Hawaiian Islands. In B. M. Riegl and R. E. Dodge (eds.), Coral Reefs of the USA. Springer Science + Business Media B.V., Dordrecht, Netherlands, pp. 519–571. Rundel, P. W. 1994. Tropical alpine climates and environments. In P. W. Rundel, A. P. Smith, and F. C. Meinzer (eds.), Tropical Alpine Environments: Plant Form and Function. Cambridge University Press, Cambridge, pp. 21–44. Sharp, W. D., and D. A. Clague. 2006. 50 Ma initiation of

The references here, organized by chapter, are largely selected to include those papers published since the Manual of the Flowering Plants of Hawai‘i was published in 1990. For chapters 4 and 5, these references will provide a basis for reviewing recent taxonomic studies on individual families and genera. Papers describing new species but without a broader review are not included. Individual descriptions for new species since the flora can be found by linking to the Smithsonian Institution’s Flora of the Hawaiian Islands Web site, available at http://botany.si.edu/ pacificislandbiodiversity/hawaiianflora/index.htm. Detailed infor­ mation on species that are federally listed as endangered can be found on the Web in documents produced by the U.S. Fish and Wildlife Service in designating critical habitat and in five-year evaluations of the status of endangered species. CHAPTER 1. LANDFORMS AND CLIMATE

Carlquist, S. 1992. Hawaii: A Natural History: Geology, Climate, Native Flora and Fauna Above the Shoreline. National Tropical Botanical Garden, Lāwa‘i, Kaua‘i (reprint of the 1970 edition). Carson, H. L., and D. A. Clague. 1995. Geology and biogeography of the Hawaiian Islands. In V. A. Funk and W. L. Wagner (eds.), Hawaiian Biogeography: Evolution on a Hot Spot Archipelago. Smithsonian Institution Press, Washington, D.C., pp. 14–29. Clague, D. A., J. C. Braga, D. Bassi, P. D. Fullagar, W. Renema, and J. M. Webster. 2010. The maximum age of Hawaiian terrestrial lineages: Geological constraints from Koko Seamount. Journal of Biogeography 37:1022–1033. Fleischer, R. C., C. E. McIntosh, and C. L. Tarr. 1998. Evolution on a volcanic conveyor belt: Using phylogeographic reconstructions and K-Ar-based ages of the Hawaiian Islands to estimate molecular evolutionary rates. Molecular Ecology 7:533–545. Giambelluca, T. W., and D. Nullet. 1992. Evaporation at high elevations in Hawaii. Journal of Hydrology 136:219–235. Giambelluca, T. W., Q. Chen, A. G. Frazier, J. P. Price, Y.-L. Chen, P.-S. Chu, J. K. Eischeid, and D. M. Delparte. 2013. Online rainfall atlas of Hawai‘i. Bull. Amer. Meteor. Soc. 94, 313– 316, doi: 10.1175/BAMS-D-11-00228.1. Available at http:// rainfall.geography.hawaii.edu. Hazlett, R. W., and D. W. Hyndman. 2003. Roadside Geology of Hawai‘i. Mountain Press, Missoula, MT.

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Hawaiian-­Emperor bend records major change in Pacific plate motion. Science 313:1281–1284. Ziegler, A. C. 2005. Hawaiian Natural History, Ecology, and Evolution. University of Hawai‘i Press, Honolulu. CHAPTER 2. PLANT DIVERSITY AND EVOLUTION

Baldwin, B. G., D. J. Crawford, J. Francisco-Ortega, S.-C. Kim, T. Sang, and T. F. Stuessy. 1998. Molecular phylogenetic insights on the origin and evolution of oceanic island plants. In D. E. Soltis, P. Soltis, and J. J. Doyle (eds.), Molecular Systematics of Plants, II: DNA sequencing. Kluwer Academic Publishers, pp. 410–441. Baldwin, B. G., and W. L. Wagner. 2010. Hawaiian angiosperm radiations of North American origin. Annals of Botany 105:849–879. Carlquist, S. 1974. Island Biology. Columbia University Press, New York. ———. 1992. Hawaii: A Natural History: Geology, Climate, Native Flora and Fauna Above the Shoreline. National Tropical Botanical Garden, Lāwa‘i, Kaua‘i (reprint of the 1970 edition). Carr, G. D. 1998. Chromosome evolution and speciation in Hawaiian flowering plants. In T. F. Stuessy and M. Ono (eds.), Evolution and Speciation of Island Plants. Cambridge University Press, New York, pp. 5–47. Castro, S. A., C. C. Daehler, L. Silva, C. W. Torres-Santana, J. A. Reyes-Betancort, R. Atkinson, P. Jaramillo, A. Guezou, and F. M. Jaksic. 2010. Floristic homogenization as a teleconnected trend in oceanic islands. Diversity and Distributions 16:902–910. Cowie, R. H., and B. S. Holland. 2006. Dispersal is fundamental to biogeography and the evolution of biodiversity on oceanic islands. Journal of Biogeography 33:193–198. ———. 2008. Molecular biogeography and diversification of the endemic terrestrial fauna of the Hawaiian Islands. Philosophical Transactions of the Royal Society of London, B. Biological Sciences 363:3363–3376. Craddock, E. M. 2000. Speciation processes in the adaptive radiation of Hawaiian plants and animals. Evolutionary Biology 31:1–53. Dominguez Lozano, F., J. Price, R. Otto, and J. Maria Fernandez-Palacios. 2010. Using taxonomic and phylogenetic evenness to compare diversification in two island floras. Perspectives in Plant Ecology Evolution and Systematics 12:93–106. Eldon, J., J. P. Price, K. Magnacca, and D. K. Price. 2013. Patterns and processes in complex landscapes: testing alternative biogeographical hypotheses through integrated analysis of phylogeography and community ecology in Hawai‘i. Molecular Ecology 22:3613–3628. Elix, J. A., and P. M. McCarthy. 1998. Catalogue of the lichens of the smaller Pacific islands. Bibliotheca Lichenologica 70: 1–361. Ellstrand, N. C., R. Whitkus, and L. H. Rieseberg. 1996. Distribution of spontaneous plant hybrids. Proceedings of the National Academy of Sciences 93:5090–5093. Funk, V. A., and W. L. Wagner. 1995. Biogeography of seven ancient Hawaiian plant lineages. In V. A. Funk and W. L.

Wagner (eds.), Hawaiian Biogeography: Evolution on a Hot Spot Archipelago. Smithsonian Institution Press, Washington, D.C. Funk, V. A., and W. L. Wagner (eds.). 1995. Hawaiian Biogeography: Evolution on a Hot Spot Archipelago. Smithsonian Institution Press, Washington, D.C. Gillespie, R. G. 2002. Biogeography of spiders on remote oceanic islands of the Pacific: Archipelagoes as stepping stones? Journal of Biogeography 29:655–662. Givnish T. J., K. J. Sytsma, J. F. Smith, and W. J. Hahn. 1994. Thorn-like prickles and heterophylly in Cyanea: Adaptations to extinct avian browsers on Hawaii. Proceedings of the National Academy of Sciences 91:2810–2814. Gruner, D. S., N. J. Gotelli, J. P. Price, and R. H. Cowie. 2008. Does species richness drive speciation? A reassessment with the Hawaiian biota. Ecography 31:279–285. Heads, M. 2012. Molecular Panbiogeography of the Tropics. University of California Press, Berkeley. Jorgensen, S., and R. Mauricio. 2005. Hybridization as a source of evolutionary novelty: Leaf shape in a Hawaiian composite. Genetica 123:171–179. Lozano, F. D., J. Price, R. Otto, and J. M. Fernandez-Palacios. Using taxonomic and phylogenetic evenness to compare diversification in two island floras. Perspectives in Plant Ecology Evolution and Systematics 12:93–106. Palmer, D. D. 2008. Hawai‘i’s Ferns and Fern Allies. University of Hawai‘i Press, Honolulu. Price, J. P. 2004. Floristic biogeography of the Hawaiian Islands: Influences of area, environment and paleogeography. Journal of Biogeography 31:487–500. Price, J. P., and D. A. Clague. 2002. How old is the Hawaiian biota? Geology and phylogeny suggest recent divergence. Proceedings of the Royal Society of London Series B: Biological Sciences 269:2429–2435. Price, J. P., and W. L. Wagner. 2011. A phylogenetic basis for species-area relationships among three Pacific Island floras. American Journal of Botany 98:449–459. Sadler, J. P. 1999. Biodiversity on oceanic islands: A palaeoecological assessment. Journal of Biogeography 26:75–87. Sakai, A. K., W. L. Wagner, D. M. Ferguson, and D. R. Herbst. 1995. Biogeographical and ecological correlates of dioecy in the Hawaiian flora. Ecology 76:2530–2543. ———. 1995. Origins of dioecy in the Hawaiian flora. Ecology 76:2517–2529. Schultz, S. T., and F. R. Ganders. 1996. Evolution of unisexuality in the Hawaiian flora: A test of microevolutionary theory. Evolution 50:842–855. Smith, C. W. 2006. Checklist of lichens and lichenicolous fungi of Hawai‘i (USA). Available at http://www.biologie.uni-​ hamburg.de/checklists/oceania/usa_hawaii_l.htm. Sohmer, S. H., and R. Gustafson.1987. Plants and Flowers of Hawai‘i. University of Hawai‘i Press, Honolulu. Staples G. W., and C. T. Imada. 2006. Checklist of Hawaiian Anthocerotes and hepatics. Tropical Bryology 28:15–47. Staples, G. W., C. T. Imada, W. J. Hoe, and C. W. Smith. 2004. A revised checklist of Hawaiian mosses. Tropical Bryology 25:35–69. Wagner, W. L., D. R. Herbst, and S. H. Sohmer (eds.). 1990. Manual of the Flowering Plants of Hawai‘i (2 vols.). University

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of Hawai‘i Press, Honolulu (republished in 1999 with a supplement). Whittaker, R. J., K. A. Triantis, and R. J. Ladle. 2008. A general dynamic theory of oceanic island biogeography. Journal of Biogeography 35:977–994. CHAPTER 3. PLANT COMMUNITIES

Allen, M. S., and G. M. Murakami. 1999. Lana‘i Island’s arid lowland vegetation in late prehistory. Pacific Science 53:88–112. Athens, J. S. 1997. Hawaiian native lowland vegetation in prehistory. In P. V. Kirch and T. L. Hunt (eds.), Historical Ecology in the Pacific Islands: Prehistoric Environmental and Landscape Change. Yale University Press, New Haven, CT, pp. 248–270. Athens, J. S., J. V. Ward, and D. W. Blinn. 2007. Vegetation history of Laysan Island, Northwestern Hawaiian Islands. Pacific Science 61:17–37. Boehmer, H. J., H. H. Wagner, J. D. Jacobi, G. C. Gerrish, and D. Mueller-Dombois. 2013. Rebuilding after collapse: Evidence for long-term cohort dynamics in the native Hawaiian rain forest. Journal of Vegetation Science 24:639–650. Brauman, K. A., D. L. Freyberg, and G. C. Daily. 2010. Forest structure influences on rainfall partitioning and cloud interception: A comparison of native forest sites in Kona, Hawai‘i. Agricultural and Forest Meteorology 150:265–275. Clarkson, B. D. 1998. Vegetation succession (1967–89) on five recent montane lava flows, Mauna Loa, Hawaii. New Zealand Journal of Ecology 22:1–9. Crausbay, S. D., and S. C. Hotchkiss. Strong relationships between vegetation and two perpendicular climate gradients high on a tropical mountain in Hawai‘i. Journal of Biogeography 37:1160–1174. Gagné, W. C., and L. W. Cuddihy. 1990. Vegetation. In W. L. Wagner, D. R. Herbst, and S. H. Sohmer (eds.), Manual of the Flowering Plants of Hawai‘i. University of Hawai‘i Press, Honolulu (republished in 1999 with a supplement), pp. 45–116. Gillespie, T. W., G. Keppel, S. Pau, J. P. Price, T. Jaffré, J-Y. Meyer, and K. O’Neill. 2011. Floristic composition and natural history characteristics of dry forests in the Pacific. Pacific Science 65:127–141. Gillespie, T. W., G. Keppel, S. Pau, J. P. Price, T. Jaffré, K. O’Neill, and M. Huston. 2013. Scaling species richness and endemism of tropical dry forests on oceanic islands. Diversity and Distributions 19:896–906. Gon, S. M. 1994. A Hawaiian bog in the Ko‘olau Mountains of O‘ahu? Evidence from community structure and diagnostic species. Newsletter of the Hawaiian Botanical Society 33:89–96. Hart, P. J. 2010. Tree growth and age in an ancient Hawaiian wet forest: Vegetation dynamics at two spatial scales. Journal of Tropical Ecology 26:1–11. Hatfield, J. S., W. A. Link, D. K. Dawson, and E. L. Lindquist. 1996. Coexistence and community structure of tropical trees in a Hawaiian montane rain forest. Biotropica 28:746–758. Herbert, D. A., J. H. Fownes, and P. M. Vitousek. 1999. Hurricane damage to a Hawaiian forest: Nutrient supply rate affects resistance and resilience. Ecology 80:908–920.

Hoof, J., L. Sack, D. T. Webb, and E. T. Nilsen. 2008. Contrasting structure and function of pubescent and glabrous varieties of Hawaiian Metrosideros polymorpha (Myrtaceae) at high elevation. Biotropica 40:113–118. Hotchkiss, S., and J. O. Juvik. 1999. A Late-Quaternary pollen record from Ka‘au Crater, O‘ahu, Hawai‘i. Quaternary Research 52:115–128. Juvik, J. O., B. T. Rodomsky, J. P. Price, E. W. Hansen, and C. Kueffer. 2011. The upper limits of vegetation on Mauna Loa, Hawaii: A 50th-anniversary reassessment. Ecology 92:518–525. Kitayama, K., and D. Mueller-Dombois. 1994. An altitudinal transect analysis of the windward vegetation on Haleakala, a Hawaiian island mountain: (2) Vegetation zonation. Phytocoenologia 24:135–154. Kellner, J. R., and G. P. Asner. 2009. Convergent structural responses of tropical forests to diverse disturbance regimes. Ecology Letters 12:887–897. Kolivras, K. N., and A. C. Comrie. 2007. Regionalization and variability of precipitation in Hawaii. Physical Geography 28:76–96. Leuschner, C. 1996. Timberline and alpine vegetation on the tropical and warm-temperate oceanic islands of the world: Elevation, structure and floristics. Vegetatio 123:193–206. Loope, L. L. 2000. Vegetation of the Hawaiian Islands. In M. G. Barbour and W. D. Billings (eds.), North American Terrestrial Vegetation, 2nd ed. Cambridge University Press, Cambridge, 661–688. Loope, L. L., A. C. Medeiros, and B. H. Gagné. 1991. Studies in the montane bogs of Haleakala National Park: Aspects of the history and biology of the montane bogs. University of Hawai‘i Cooperative National Park Resources Studies Unit Technical Report No. 76:1–43. Mueller-Dombois, D. 1987. Forest dynamics in Hawaii. Trends in Ecology & Evolution 2:216–220. ———. 2002. Forest vegetation across the tropical Pacific: A biogeographically complex region with many analogous environments. Plant Ecology 163:155–176. ———. 2006. Long-term rain forest succession and landscape change in Hawai‘i: The “Maui Forest Trouble” revisited. Journal of Vegetation Science 17:685–692. ———. 2000. Rain forest establishment and succession in the Hawaiian Islands. Landscape and Urban Planning 51:147–157. Mueller-Dombois, D., and F. R. Fosberg. 1998. Vegetation of the Tropical Pacific Islands. Springer Verlag, Berlin. Noguchi, Y. 1992. Hydrometeorological differences between opposite valley slopes and vegetation asymmetry in Hawaii. Journal of Vegetation Science 3:231–238. ———. 1992. Vegetation asymmetry in Hawaii under the tradewind regime. Journal of Vegetation Science 3:223–230. Pau, S., T. W. Gillespie, and J. P. Price. 2009. Natural history, biogeography, and endangerment of Hawaiian dry forest trees. Biodiversity and Conservation 18:3167–3182. Restrepo, C., and P. Vitousek. 2001. Landslides, alien species, and the diversity of a Hawaiian montane mesic ecosystem. Biotropica 33:409–420. Stemmermann, L., and T. Ihsle. 1993. Replacement of Metrosideros polymorpha, ohia, in Hawaiian dry forest succession. Biotropica 25:36–45.

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Vitousek, P., G. P. Asner, O. A. Chadwick, and S. Hotchkiss. 2009. Landscape-level variation in forest structure and biogeochemistry across a substrate age gradient in Hawaii. Ecology 90:3074–3086. Vogl, R. J., and J. Henrickson. 1971. Vegetation of an alpine bog on East Maui, Hawaii. Pacific Science 25:475–483. Warren, S. D. 2004. Degradation and recovery of vegetation on Kaho‘olawe Island, Hawaii: A photographic journey. Pacific Science 58:461–495. Zimmerman, N., R. F. Hughes, S. Cordell, P. Hart, H. K. Chang, D. Perez, R. K. Like, and R. Ostertag. 2008. Patterns of primary succession of native and introduced plants in lowland wet forests in eastern Hawai‘i. Biotropica 40:277–284. CHAPTER 4. FLOWERING PLANTS

Angiosperm Phylogeny Group. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 161:105–121. Mabberley, D. J. 2008. Mabberley’s Plant-Book: A Portable Dictionary of Plants, Their Classifications and Uses (3rd edition). Cambridge University Press, Cambridge. Morden, C. W., V. Caraway, and T. J. Motley. 1996. Development of a DNA library for native Hawaiian plants. Pacific Science 50:324–335. Schonenberger, J., A. A. Anderberg, and K. J. Sytsma. 2005. Molecular phylogenetics and patterns of floral evolution in the Ericales. International Journal of Plant Sciences 166:265–288. Wagner, W. L., D. R. Herbst, and S. H. Sohmer (eds.). 1990. Manual of the Flowering Plants of Hawai‘i. University of Hawai‘i Press, Honolulu (republished in 1999 with a supplement). Wood, K. R. 2007. New plant records, rediscoveries, range extensions, and possible extinctions within the Hawaiian Islands. Bishop Museum Occasional Papers 96:13–17. ———. 2012. Possible extinctions, rediscoveries, and new plant records within the Hawaiian Islands. In N. L. Evenhuis and L. G. Eldredge (eds.), Records of the Hawaii Biological Survey for 2011. Bishop Museum Occasional Papers 113:91–102. DICOTS

Amaranthaceae Eliasson, U. H. 2004. The evolutionary patterns of the plant family Amaranthaceae on the Galapagos and Hawaiian Islands. Journal of the Torrey Botanical Society 131:105–109. Muller, K., and T. Borsch. 2005. Phylogenetics of Amaranthaceae based on matK/trnK sequence data: Evidence from Parsimony, likelihood, and Bayesian analyses. Annals of the Missouri Botanical Garden 92:66–102. Anacardiaceae Yi, T. S., A. J. Miller, and J. Wen. 2004. Phylogenetic and biogeographic diversification of Rhus (Anacardiaceae) in the Northern Hemisphere. Molecular Phylogenetics and Evolution 33:861–879.

Apiaceae Vargas, P., B. G. Baldwin, and L. Constance. 1998. Nuclear ribosomal DNA evidence for a western North American origin of Hawaiian and South American species of Sanicula (Apiaceae). Proceedings of the National Academy of Sciences 95:235–240. ———. 1997. Phylogenetic insights on historical biogeography in Sanicula (Apiaceae) from ITS sequence data: A western North American origin of the Hawaiian sanicles. American Journal of Botany 84: 239–240. Apocynaceae Middleton, D. J. 2002. Revision of Alyxia (Apocynaceae). Part 2: Pacific Islands and Australia. Blumea 47:1–93. Aquifoliaceae Cuenoud, P., M. A. D. Martinez, P. A. Loizeau, R. Spichiger, S. Andrews, and J. F. Manen. 2000. Molecular phylogeny and biogeography of the genus Ilex L. (Aquifoliaceae). Annals of Botany 85:111–122. Araliaceae Costello, A., and T. J. Motley. 2004. The development of the superior ovary in Tetraplasandra (Araliaceae). American Journal of Botany 91:644–655. ———. 2001. Molecular systematics of Tetraplasandra, Munroidendron and Reynoldsia sandwicensis (Araliaceae) and the evolution of superior ovaries in Tetraplasandra. Edinburgh Journal of Botany 58:229–242. ———. 2007. Phylogenetics of the Tetraplasandra group (Araliaceae) inferred from ITS, 5S-NTS, and morphology. Systematic Botany 32:464–477. Lowry, P. P., and G. M. Plunkett. 2010. Recircumscription of Polyscias (Araliaceae) to include six related genera, with a new infrageneric classification and a synopsis of species. Plant Diversity and Evolution 128:55–84. Mitchell, A., R. Li, J. W. Brown, I. Schönberger, and J. Wen . 2012. Ancient divergence and biogeography of Raukaua (Araliaceae) and close relatives in the southern hemisphere. Australian Systematic Botany 25:432–446. Plunkett, G. M., and P. P. Lowry. 2010. Paraphyly and polyphyly in Polyscias sensu lato: Molecular evidence and the case for recircumscribing the “pinnate genera” of Araliaceae. Plant Diversity and Evolution 128:23–54. Asteraceae Baldwin, B. G. 2000. Adaptive radiation of the Hawaiian silversword alliance: Congruence and conflict of phylogenetic evidence from molecular and non-molecular investigations. In T. J. Givnish and K. J. Sytsma (eds.), Molecular Evolution and Adaptive Radiation. Cambridge University Press, Cambridge, pp. 103–128. ———. 2007. Adaptive radiation of shrubby tarweeds (Deinandra) in the California Islands parallels diversification of the Hawaiian silversword alliance (Compositae-Madiinae). American Journal of Botany 94:237–248. ———. 2006. Contrasting patterns and processes of evolutionary change in the tarweed-silversword lineage: Revisiting

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Clausen, Keck, and Hiesey’s findings. Annals of the Missouri Botanical Garden 93:64–93. ———. 2003. Natural history of the continental tarweeds and the Hawaiian silversword alliance. In S. Carlquist, B. G. Baldwin, and G. D. Carr (eds.), Tarweeds and Silverswords: Evolution of the Madiinae. Missouri Botanical Garden Press, St. Louis, pp. 1–16. ———. 2003. A phylogenetic perspective on the origin and evolution of Madiinae. In S. Carlquist, B. G. Baldwin, and G. Carr (eds.), Tarweeds and Silverswords: Evolution of the Madiinae. Missouri Botanical Garden Press, St. Louis, pp. 193–228. Baldwin, B. G., and E. A. Friar. 2010. Dubautia carrii and D. hanaulaensis, new species of the Hawaiian silversword alliance (Compositae, Madiinae) from Moloka‘i and Maui. Novon 20:1–8. Baldwin, B. G., and R. H. Robichaux. 1995. Historical biogeography and ecology of the Hawaiian silversword alliance (Asteraceae). In W. L. Wagner and V. A. Funk (eds.), Hawaiian Biogeography: Evolution on a Hot Spot Archipelago. Smithsonian Institution Press, Washington, D.C., pp. 259–287. Baldwin, B. G., and M. J. Sanderson. 1998. Age and rate of diversification of the Hawaiian silversword alliance (Compositae). Proceedings of the National Academy of Sciences 95:9402–9406. Baldwin, B. G., and B. L. Wessa. 2000. Origin and relationships of the tarweed-silversword lineage (Compositae-Madiinae). American Journal of Botany 87:1890–1908. Carlquist, S. 2003. Leaf anatomy of Madiinae and adaptive radiation. In S. Carlquist, B. G. Baldwin, and G. D. Carr (eds.), Tarweeds and Silverswords: Evolution of the Madiinae (Asteraceae). Missouri Botanical Garden Press, St. Louis. Carr, G. D. 1999. A reassessment of Dubautia (Asteraceae: Heliantheae-Madiinae) on Kaua‘i. Pacific Science 53:144–158. Friar, E. A., J. M. Cruse-Sanders, and M. E. McGlaughlin. 2007. Gene flow in Dubautia arborea and D. ciliolata: The roles of ecology and isolation by distance in maintaining species boundaries despite ongoing hybridization. Molecular Ecology 16:4028–4038. Friar, E. A., T. Ladoux, E. H. Roalson, and R. H. Robichaux. 2000. Microsatellite analysis of a population crash and bottleneck in the Mauna Kea silversword, Argyroxiphium sandwicense ssp. sandwicense (Asteraceae), and its implications for reintroduction. Molecular Ecology 9:2027–2034. Friar, E. A., L. M. Prince, J. M. Cruse-Sanders, M. E. McGlaughlin, C. A. Butterworth, and B. G. Baldwin. 2008. Hybrid origin and genomic mosaicism of Dubautia scabra (Hawaiian Silversword Alliance; Asteraceae, Madiinae). Systematic Botany 33:589–597. Friar, E. A., L. M. Prince, E. H. Roalson, M. E. McGlaughlin, J. M. Cruse-Sanders, S. J. De Groot, and J. M. Porter. 2006. Ecological speciation in the East Maui-endemic Dubautia (Asteraceae) species. Evolution 60:1777–1792. Friar, E. A., and R. H. Robichaux. 2003. Conservation genetics of the silversword alliance. In S. Carlquist, B. G. Baldwin, and G. D. Carr (eds.), Tarweeds and Silverswords: Evolution of the Madiinae (Asteraceae). Missouri Botanical Garden Press, St. Louis.

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FURTHER READING

Lamiaceae Lindqvist, C., and V. A. Albert. 2002. Origin of the Hawaiian endemic mints within North American Stachys (Lamiaceae). American Journal of Botany 89:1709–1724. Lindqvist, C., T. J. Motley, J. J. Jeffrey, and V. A. Albert. 2003. Cladogenesis and reticulation in the Hawaiian endemic mints (Lamiaceae). Cladistics: The International Journal of the Willi Hennig Society 19:480–495. Morden, C. W., and W. Loeffler. 1999. Fragmentation and genetic differentiation among subpopulations of the endangered Hawaiian mint Haplostachys haplostachya (Lamiaceae). Molecular Ecology 8:617–625. Roy, T., T.-H. Chang, T. Lan, and C. Lindqvist. 2013. Phylogeny and biogeography of New World Stachydeae (Lamiaceae) with emphasis on the origin and diversification of Hawaiian and South American taxa. Molecular Phylogenetics and Evolution 69:218–238. Wagner, W. L. 1999. A new species of Hawaiian Phyllostegia (Lamiaceae) from Kaua‘i and recognition of a Wai‘anae Mountain, O‘ahu, endangered variety of Phyllostegia parviflora. Novon 9:280–283. Loganiaceae Motley, T. J., and G. D. Carr. 1998. Artificial hybridization in the Hawaiian endemic genus Labordia (Loganiaceae). American Journal of Botany 85:654–660. Malvaceae Dejoode, D. R., and J. F. Wendel. 1992. Genetic diversity and origin of the Hawaiian Islands cotton, Gossypium tomentosum. American Journal of Botany 79:1311–1319. Funk, V. A., and W. L. Wagner. 1995. Biogeography of seven ancient Hawaiian plant lineages. In V. A. Funk and W. L. Wagner (eds.), Hawaiian Biogeography: Evolution on a Hot Spot Archipelago. Smithsonian Institution Press, Washington, D.C., pp. 160–194. Seelanan T., A. Schnabel, and J. F. Wendel. 1997. Congruence and consensus in the cotton tribe (Malvaceae). Systematic Botany 22:259–290. Myrtaceae Aradhya, K. M., D. Mueller-Dombois, and T. A. Ranker. 1991. Genetic-evidence for recent and incipient speciation in the evolution of Hawaiian Metrosideros (Myrtaceae). Heredity 67:129–138. ———. 1993. Genetic-structure and differentiation in Metrosideros polymorpha (Myrtaceae) along altitudinal gradients in Maui, Hawaii. Genetical Research 61:159–170. Crawford, N. G., C. Hagen, H. F. Sahli, E. A. Stacy, and T. C. Glenn. 2008. Fifteen polymorphic microsatellite DNA loci from Hawaii’s Metrosideros polymorpha (Myrtaceae: Myrtales), a model species for ecology and evolution. Molecular Ecology Resources 8:308–310. Harbaugh, D. T., W. L. Wagner, D. M. Percy, H. F. James, and R. C. Fleischer. 2009. Genetic structure of the polymorphic Metrosideros (Myrtaceae) complex in the Hawaiian Islands using nuclear microsatellite data. PLoS ONE 4(3) e4698. doi:10.1371/journal.pone.0004698.

Mueller-Dombois, D. 1994. Vegetation dynamics and the evolution of Metrosideros polymorpha in Hawaii. Phytocoenologia 24:609–614. Percy, D. M., A. M. Garver, W. L. Wagner, H. F. James, C. W. Cunningham, S. E. Miller, and R. C. Fleischer. 2008. Progressive island colonization and ancient origin of Hawaiian Metrosideros (Myrtaceae). Proceedings of the Royal Society B: Biological Sciences 275:1479–1490. Wright, M. E., and T. A. Ranker. 2010. Dispersal and habitat fidelity of bog and forest growth forms of Hawaiian Metro­ sideros (Myrtaceae). Botanical Journal of the Linnean Society 162:558–571. Wright, S. D., C. G. Yong, S. R. Wichman, J. W. Dawson, and R. C. Gardner. 2001. Stepping stones to Hawaii: A trans-equatorial dispersal pathway for Metrosideros (Myrtaceae) inferred from nrDNA (ITS plus ETS). Journal of Biogeography 28:769–774. Pentaphylacaceae Prince, L. M., and C. R. Parks. 2001. Phylogenetic relationships of Theaceae inferred from chloroplast DNA sequence data. American Journal of Botany 88:2309–2320. Phyllanthaceae Samuel, R., H. Kathriarachchi, P. Hoffmann, M. H. Barfuss, K. J. Wurdack, C. C. Davis, and M. W. Chase. 2005. Molecular phylogenetics of Phyllanthaceae: Evidence from plastid Matk and nuclear PHYC sequences. American Journal of Botany 92:132–141. Piperaceae Smith J. F., A. C. Stevens, E. J. Tepe, and C. Davidson. 2008. Placing the origin of two species-rich genera in the late Cretaceous with later species divergence in the Tertiary: A phylogenetic, biogeographic and molecular dating analysis of Piper and Peperomia (Piperaceae). Plant Systematics and Evolution 275:9–30. Pittosporaceae Bacon, C. D., G. J. Allan, E. A. Zimmer, and W. L. Wagner. 2011. Genome scans reveal high levels of gene flow in Hawaiian Pittosporum. Taxon 60:733–741. Gemmill, C. E. C., G. J. Allan, W. L. Wagner, and E. A. Zimmer. 2002. Evolution of insular Pacific Pittosporum (Pittosporaceae): Origin of the Hawaiian radiation. Molecular Phylogenetics and Evolution 22:31–42. Plantaginaceae Dunbar-Co, S., M. J. Sporck, and L. Sack. 2009. Leaf trait diversification and design in seven rare taxa of the Hawaiian Plantago radiation. International Journal of Plant Sciences 170:61–75. Dunbar-Co, S., and A. M. Wieczorek. 2011. Genetic structure among populations in the endemic Hawaiian Plantago lineage: Insights from microsatellite variation. Plant Species Biology 26:134–144. Dunbar-Co, S., A. M. Wieczorek, and C. W. Morden. 2008. Molecular phylogeny and adaptive radiation of the endemic Hawaiian Plantago species (Plantaginaceae). American Journal of Botany 95:1177–1188.

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Portulacaceae Kim, I., and G. D. Carr. 1990. Cytogenetics and hybridization of Portulaca in Hawaii. Systematic Botany 15:370–377. Primulaceae Hao G., Y. M. Yuan, C. M. Hu, X. J. Ge, and N. X. Zhao. 2004. Molecular phylogeny of Lysimachia (Myrsinaceae) based on chloroplast trnL-F and nuclear ribosomal ITS sequences. Molecular Phylogenetics and Evolution 31:323–339. Marr, K. L., and B. A. Bohm. 1997. A taxonomic revision of the endemic Hawaiian Lysimachia (Primulaceae) including three new species. Pacific Science 51:254–287. Wood, K. R. 2012. Rediscovery of Lysimachia venosa (Wawra) H. St. John on Kaua‘i, Hawaiian Islands. In N. L. Evenhuis and L. G. Eldredge (eds.), Records of the Hawaii Biological Survey for 2012. Bishop Museum Occasional Papers 114:37–38. Rhamnaceae Kwon, J. A., and C. W. Morden. 2002. Population genetic structure of two rare tree species (Colubrina oppositifolia and Alphitonia ponderosa, Rhamnaceae) from Hawaiian dry and mesic forests using random amplified polymorphic DNA markers. Molecular Ecology 11:991–1001. Rosaceae Howarth, D. G., D. E. Gardner, and C. W. Morden. 1997. Phylogeny of Rubus subgenus Idaeobatus (Rosaceae) and its implications toward colonization of the Hawaiian Islands. Systematic Botany 22:433–441. Rubiaceae Anderson, C. L., J. H. E. Rova, and L. Andersson. 2001. Molecular phylogeny of the tribe Anthospermeae (Rubiaceae): Systematic and biogeographic implications. Australian Systematic Botany 14:231–244. Heads, M. J. 1996. Biogeography, taxonomy and evolution in the Pacific genus Coprosma (Rubiaceae). Candollea 51:381–405. Motley, T. J., L. Struwe, and V. A. Albert. 1998. Molecular systematics of Hawaiian Hedyotis (Rubiaceae). American Journal of Botany 85:146. Nepokroeff, M., K. J. Sytsma, W. L. Wagner, and E. A. Zimmer. 2003. Reconstructing ancestral patterns of colonization and dispersal in the Hawaiian understory shrub genus Psychotria (Rubiaceae): A comparison of parsimony and likelihood approaches. Systematic Biology 52:820–838. Razafimandimbison, S. G., and B. Bremer. 2011. Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae). Adansonia, sér. 3, 33:283–309. Terrell, E. E., H. E. Robinson, W. L. Wagner, and D. H. Lorence. 2005. Resurrection of genus Kadua for Hawaiian Hedyotidinae (Rubiaceae), with emphasis on seed and fruit characters and notes on South Pacific species. Systematic Botany 30:818–833 Rutaceae Funk, V. A., and W. L. Wagner. 1995. Biogeography of seven ancient Hawaiian plant lineages. In V. A. Funk and W. L.

Wagner (eds.), Hawaiian Biogeography: Evolution on a Hot Spot Archipelago. Smithsonian Institution Press, Washington, D.C., pp. 160–194. Harbaugh, D. T., W. L. Wagner, G. J. Allan, and E. A. Zimmer. 2009. The Hawaiian Archipelago is a stepping stone for dispersal in the Pacific: An example from the plant genus Melicope (Rutaceae). Journal of Biogeography 36:230–241. Hartley, T. G. 2001. On the taxonomy and biogeography of Euodia and Melicope (Rutaceae). Allertonia 8:1–319. Santalaceae Harbaugh, D. T. 2008. Polyploid and hybrid origins of Pacific Island sandalwoods (Santalum, Santalaceae) inferred from low-copy nuclear and flow cytometry data. International Journal of Plant Sciences 169:677–685. Harbaugh, D. T., and B. G. Baldwin. 2007. Phylogeny and biogeography of the sandalwoods (Santalum, Santalaceae): Repeated dispersals throughout the Pacific. American Journal of Botany 94:1028–1040. Harbaugh, D. T., H. L. Oppenheimer, K. R. Wood, and W. L. Wagner. 2010. Taxonomic revision of the endangered Hawaiian red-flowered sandalwoods (Santalum) and discovery of an ancient hybrid species. Systematic Botany 35:827–838. Molvray, M., P. J. Kores, and M. W. Chase. 1999. Phylogenetic relationships within Korthalsella (Viscaceae) based on nuclear ITS and plastid trnL-F sequence data. American Journal of Botany 86:249–260. Sapotaceae Smedmark, J. E. E., and A. A. Anderberg. 2007. Boreotropical migration explains hybridization between geographically distant lineages in the pantropical clade Sideroxyleae (Sapotaceae). American Journal of Botany 94:1491–1505. Swenson, U., and A. A. Anderberg. 2005. Phylogeny, character evolution, and classification of Sapotaceae (Ericales). Cladistics 21:101–130. Swenson, U., J. Munzinger, and I. V. Bartish. 2007. Molecular phylogeny of Planchonella (Sapotaceae) and eight new species from New Caledonia. Taxon 56:329–354. Thymelaeaceae Mayer, S. S. 1991. Morphological variation in Hawaiian Wikstroemia (Thymelaeaceae). Systematic Botany 16:693–704. Mayer, S. S., and D. Charlesworth. 1992. Genetic-evidence for multiple origins of dioecy in the Hawaiian shrub Wikstroemia (Thymelaeaceae). Evolution 46:207–215. Urticaceae Loeffler, W. F., and C. W. Morden. 2003. Genetic diversity and biogeography of the Hawaiian cordage plant, olona (Touchardia latifolia; Urticaceae), based on RAPD markers. Biochemical Systematics and Ecology 31:1323–1335. Violaceae Ballard, H. E., Jr., and K. J. Sytsma. 2000a. Evolution and biogeography of the woody Hawaiian violets (Viola, Violaceae): Arctic origins, herbaceous ancestry and bird dispersal. Evolution 54:1521–1532.

FURTHER READING

Havran, J. C., K. J. Sytsma, and H. E. Ballard, Jr. 2009. Evolutionary relationships, interisland biogeography, and molecular evolution in the Hawaiian violets (Viola: Violaceae). American Journal of Botany 96:2087–2099.

Xanthorrhoeaceae

MONOCOTS

Chase, M. W., J. L. Reveal, and M. F. Fay. 2009. A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society 161:132–136.

Arecaceae

CHAPTER 5. FERNS AND FERN ALLIES

Bacon, C. D., G. P. Johnson, H. Meimberg, P. Puppo, M. P. Simmons, and W. L. Wagner. 2011b. Development of microsatellites in the Hawaiian endemic palm Pritchardia martii (Arecaceae) and their utility in congeners. American Journal of Botany 98:E139–E140. Chapin, M. H., M. Maunder, and K. E. Horak. 2007. A preliminary study of regeneration in wild populations of threatened endemic Hawaiian palms (Pritchardia; Arecaceae). Pacific Conservation Biology 13:20–28. Chapin, M. H., K. R. Wood, S. P. Perlman, and M. Maunder. 2004. A review of the conservation status of the endemic Pritchardia palms of Hawaii. Oryx 38:273–281. Hodel, D. R. 2009. A new species of Pritchardia and the rediscovery of P. lowreyana on Oahu, Hawaii. Palms 53:173–179. Hodel, D. R. 2007. A review of the genus Pritchardia. Palms 51:S1–S52. Kissling, W. D., W. L. Eiserhardt, W. J. Baker, F. Borchsenius, T. L. P. Couvreur, H. Balslev, and J.-C. Svenning. 2012. Cenozoic imprints on the phylogenetic structure of palm species assemblages worldwide. Proceedings of the National Academy of Sciences 109:7379–7384. Asparagaceae Chase, M. W., J. L. Reveal, and M. F. Fay. 2009. A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society 161:132–136. Cyperaceae Chacon, J., S. Madrina, M. W. Chase, and J. J. Bruhl. 2006. Molecular phylogenetics of Oreobolus (Cyperaceae) and the origin and diversification of the American species. Taxon 55:359–366 Poaceae Carino, D. A., and C. C. Daehler 1999. Genetic variation in an apomictic grass, Heteropogon contortus, in the Hawaiian Islands. Molecular Ecology 8:2127–2132. Chemisquy, A. M., L. M. Giussani, M. A. Scataglini, E. A. Kellogg, and O. Morrone. 2010. Phylogenetic studies favour the unification of Pennisetum, Cenchrus and Odontelytrum (Poaceae): A combined nuclear, plastid and morphological analysis, and nomenclatural combinations in Cenchrus. Annals of Botany 106:107–130. Edwards, E. J., and C. J. Still. 2008. Climate, phylogeny and the ecological distribution of C 4 grasses. Ecology Letters 11:266–276. Kellogg, E. A., S. S. Aliscioni, O. Morrone, J. Pensiero, and F. Zuloaga. 2009. A phylogeny of Setaria (Poaceae, Panicoideae, Paniceae) and related genera based on the chloroplast gene ndhF. International Journal of Plant Sciences 170:117–131.

Creese, C., A. Lee, and L. Sack. 2011. Drivers of morphological diversity and distribution in the Hawaiian fern flora: Trait associations with size, growth form, and environment. American Journal of Botany 98:956–966. Geiger, J. M. O., T. A. Ranker, J. M. R. Neale, and S. T. Klimas. 2007. Molecular biogeography and origins of the Hawaiian fern flora. Brittonia 59:142–158. Palmer, D. D. 2008. Hawai‘i’s Ferns and Fern Allies. University of Hawai‘i Press, Honolulu. Smith, A. R., K. M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, and P. G. Wolf. 2006. A classification for extant ferns. Taxon 55:705–731. Snow, N., T. Ranker, and D. H. Lorence. 2012. Taxonomic changes in Hawaiian ferns and lycophytes. Bishop Museum Occasional Papers 110:11–16. Wagner, W. H., Jr. 1995. Evolution of Hawaiian ferns and fern allies in relation to their conservation status. Pacific Science 49:31–41. Wagner, W. H., Jr., F. S. Wagner, and T. Flynn. 1995. Taxonomic notes on the pteridophytes of Hawaii. Contributions from the University of Michigan Herbarium 20:241–260. Wood, K. R. 2007. New plant records, rediscoveries, range extensions, and possible extinctions within the Hawaiian Islands. Bishop Museum Occasional Papers 96:13–17. ———. 2012. Possible extinctions, rediscoveries, and new plant records within the Hawaiian Islands. In N. L. Evenhuis and L. G. Eldredge (eds.), Records of the Hawaii Biological Survey for 2011. Bishop Museum Occasional Papers 113:91–102. Aspleniaceae Aguraiuja, R. 2003. Diellia mannii (D. C. Eaton) Robins. (Aspleniaceae) rediscovered in Hawai‘i. American Fern Journal 93:154–156. Aguraiuja, R., M. Moora, and M. Zobel. 2004. Population stage structure of Hawaiian endemic fern taxa of Diellia (Aspleniaceae): Implications for monitoring and regional dynamics. Canadian Journal of Botany: Revue Canadienne de Botanique 82:1438–1445. Aguraiuja, R., and K. R. Wood. 2002. The critically endangered endemic fern genus Diellia Brack. in Hawaii: Its population structure and distribution. Fern Gazette 16:330–334. Aguraiuja, R., M. Zobel, K. Zobel, and M. Moora. 2008. Conservation of the endemic fern lineage Diellia (Aspleniaceae) on the Hawaiian Islands: Can population structure indicate regional dynamics and endangering factors? Folia Geobotanica 43:3–18. Murakami, N. 1995. Systematics and evolutionary biology of the fern genus Hymenasplenium (Aspleniaceae). Journal of Plant Research 108:257–278. Ranker, T. A., S. K. Floyd, and P. G. Trapp. 1994. Multiple

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colonizations of Asplenium-adiantum-nigrum onto the Hawaiian archipelago. Evolution 48:1364–1370. Ranker, T. A., S. K. Floyd, M. D. Windham, and P. G. Trapp. 1994. Historical biogeography of Asplenium adiantum-­ nigrum (Aspleniaceae) in North America and implications for speciation theory in homosporous pteridophytes. American Journal of Botany 81:776–781. Schneider, H., T. A. Ranker, S. J. Russell, R. Cranfill, J. M. O. Geiger, R. Aguraiuja, K. R. Wood, M. Grundmann, K. Kloberdanz, and J. C. Vogel. 2005. Origin of the endemic fern genus Diellia coincides with the renewal of Hawaiian terrestrial life in the Miocene. Proceedings of the Royal Society B: Biological Sciences 272:455–460.

Microevolutionary patterns and processes of the native Hawaiian colonizing fern Odontosoria chinensis (Lindsaeaceae). Evolution 54:828–839. Marsileaceae Wester, L., J. Delay, L. Hoang, B. Iida, N. Kalodimos, and T. Wong. 2006. Population dynamics of Marsilea villosa (Marsileaceae) on O‘ahu, Hawai‘i. Pacific Science 60:385–402. Polypodiaceae

Speer, W. D. 2008. Phylogenetic and biogeographic relationships among North American and Hawaiian Pteridium aquilinum (L.) Kuhn (Dennstaedtiaceae) based on chloroplast rps4 and rps4-trnS intergenic spacer sequences. American Fern Journal 98:179–193. Thomson, J. A. 2004. Towards a taxonomic revision of Pteridium (Dennstaedtiaceae). Telopea 10:793–803.

Labiak, P. H. 2011. Stenogrammitis, a new genus of grammitid ferns segregated from Lellingeria (Polypodiaceae). Brittonia 63:139–149. Li, J. W., and C. H. Haufler. 1999. Genetic variation, breeding systems, and patterns of diversification in Hawaiian Polypodium (Polypodiaceae). Systematic Botany 24:339–355. Ranker, T. A. 1994. Evolution of high genetic-variability in the rare Hawaiian fern Adenophorus periens and implications for conservation management. Biological Conservation 70:19–24. Ranker, T. A., J. M. O. Geiger, S. C. Kennedy, A. R. Smith, C. H. Haufler, and B. S. Parris. 2003. Molecular phylogenetics and evolution of the endemic Hawaiian genus Adenophorus (Grammitidaceae). Molecular Phylogenetics and Evolution 26:337–347. Schneider, H., A. R. Smith, R. Cranfill, T. J. Hildebrand, C. H. Haufler, and T. A. Ranker. 2004. Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae & Grammitidaceae): Exploring aspects of the diversification of epiphytic plants. Molecular Phylogenetics and Evolution 31:1041–1063.

Dryopteridaceae

Pteridaceae

Driscoll, H. E., and D. S. Barrington. 2007. Origin of Hawaiian Polystichum (Dryopteridaceae) in the context of a world phylogeny. American Journal of Botany 94:1413–1424. Geiger, J. M. O., and T A. Ranker. 2005. Molecular phylogenetics and historical biogeography of Hawaiian Dryopteris (Dryopteridaceae). Molecular Phylogenetics and Evolution 34:392–407.

Yesilyurt, J. C. 2005. The fern genus Doryopteris (Cheilanthoideae-Pteridaceae) in the Hawaiian Islands. Kew Bulletin 60:547–558.

Blechnaceae Palmer, D. D. 1996. A revision of the genus Sadleria (Blechnaceae). Pacific Science 51:288–305. Cibotiaceae Motley, T. J., and C. W. Morden. 2001. Utility of RAPD markers in evaluating the status of the Hawaiian tree fern Cibotium x heleniae. Pacific Science 55:145–155. Dennstaedtiaceae

Gleicheniaceae Russell, A. E., J. W. Raich, and P. M. Vitousek. 1998. The ecology of the climbing fern Dicranopteris linearis on windward Mauna Loa, Hawaii. Journal of Ecology 86:765–779. Hymenophyllaceae Ebihara, A., J. H. Nitta, and K. Iwatsuki. 2011. The Hymenophyllaceae of the Pacific Area. 2. Hymenophyllum (excluding subgen. Hymenophyllum). Bulletin of the National Museum of Nature and Science, Series B: Botany 36:43–59. Isoetaceae Taylor, W. C., W. H. Wagner, R. W. Hobdy, and F. R. Warshauer. 1993. Isoetes hawaiiensis: A previously undescribed quillwort from Hawaii. American Fern Journal 83:67–70. Lindsaeaceae Ranker, T. A., C. E. C. Gemmill, and P. G. Trapp. 2000.

Thelypteridaceae Medeiros, A. C., R. W. Hobdy, and W. H. Wagner. 1994. Notes on the rediscovery, status, and ecology of the very rare Hawaiian fern Christella boydiae (Thelypteridaceae). American Fern Journal 83:86–89. CHAPTER 6. POLYNESIAN INTRODUCTIONS

Athens, J. S., H. D. Tuggle, J. V. Ward, and D. J. Welch. 2002. Avifaunal extinctions, vegetation change, and Polynesian impacts in prehistoric Hawai‘i. Archaeology in Oceania 37:57–78. Bevacqua, R. F. 1994. Origin of horticulture in Southeast Asia and the dispersal of domesticated plants to the Pacific Islands by Polynesian voyagers: The Hawaiian Islands casestudy. Hortscience 29:1226–1229. Burney, D. A. 1997. Tropical islands as paleoecological laboratories: Gauging the consequences of human arrival. Human Ecology 25:437–457. Burney, D. A., and W. K. P. Kikuchi. 2006. A millennium of human activity at Makauwahi Cave, Maha Ulepu, Kaua‘i. Human Ecology 34:219–247.

FURTHER READING

Burney, L. P., and D. A. Burney. 2003. Charcoal stratigraphies for Kaua‘i and the timing of human arrival. Pacific Science 57:211–226. Cox, P. C., and S. A. Banack (eds.). 2003. Islands, Plants and Polynesians: An Introduction to Polynesian Ethnobotany. Dioscorides Press, Portland, OR. Horrocks, M., and R. B. Rechtman. 2009. Sweet potato (Ipomoea batatas) and banana (Musa sp.) microfossils in deposits from the Kona Field System, Island of Hawaii. Journal of Archaeological Science 36:1115–1126. Kirch, P. V. 2007. Three islands and an archipelago: Reciprocal interactions between humans and island ecosystems in Polynesia. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 98:85–99. Kirch, P. V., J. Coil, A. S. Hartshorn, M. Jeraj, P. M. Vitousek, and O. A. Chadwick. 2005. Intensive dryland farming on the leeward slopes of Haleakala, Maui, Hawaiian Islands: Archaeological, archaeobotanical, and geochemical perspectives. World Archaeology 37:240–258. Kirch, P. V., A. S. Hartshorn, O. A. Chadwick, P. M. Vitousek, D. R. Sherrod, J. Coil, L. Holm, and W. D. Sharp. 2004. Environment, agriculture, and settlement patterns in a marginal Polynesian landscape. Proceedings of the National Academy of Sciences 101:9936–9941. Kirch, P. V., and J. G. Kahn. 2007. Advances in Polynesian prehistory: A review and assessment of the past decade (1993– 2004). Journal of Archaeological Research 15:191–238. Krauss, B. H. 1993. Plants in Hawaiian Culture. University of Hawai‘i Press, Honolulu. Lebot, V., E. Johnston, Q. Y. Zheng, D. McKern, and D. J. Mc­ Kenna. 1999. Morphological, phytochemical, and genetic variation in Hawaiian cultivars of ‘awa (kava, Piper methysticum, Piperaceae). Economic Botany 53:407–418. Lebot, V., B. A. Meilleur, and R. M. Manshardt. 1994. Genetic diversity in eastern Polynesian Eumusa bananas. Pacific Science 48:16–31. Matisoo-Smith, E., R. M. Roberts, G. J. Irwin, J. S. Allen, D. Penny, and D. M. Lambert. 1998. Patterns of prehistoric human mobility in Polynesia indicated by mtDNA from the Pacific rat. Proceedings of the National Academy of Sciences 95:15145–15150. McCoy, M. D., and M. W. Graves. 2010. The role of agricultural innovation on Pacific Islands: A case study from Hawai‘i Island. World Archaeology 42:90–107. McCoy, M. D., M. W. Graves, and G. Murakami. 2010. Introduction of breadfruit (Artocarpus altilis) to the Hawaiian Islands. Economic Botany 64:374–381. Mueller, J. G., Y. Ogneva-Himmelberger, S. Lloyd, and J. M. Reed. Predicting prehistoric taro (Colocasia esculenta var. antiquorum) lo‘i distribution in Hawaii. Economic Botany 64:22–33. Razafimandimbison, S. G., T. D. McDowell, D. A. Halford, and B. Bremer. 2010. Origin of the pantropical and nutriceutical Morinda citrifolia L. (Rubiaceae): Comments on its distribution range and circumscription. Journal of Biogeography 37:520–529. Rieth, T. M., T. L. Hunt, C. Lipo, and J. M. Wilmshurst. 2011. The 13th century Polynesian colonization of Hawai‘i Island. Journal of Archaeological Science 38:2740–2749.

Schenck, S., M. W. Crepeau, K. K. Wu, P. H. Moore, Q. Yu, and R. Ming. 2004. Genetic diversity and relationships in native Hawaiian Saccharum officinarum sugarcane. Journal of Heredity 95:327–331. Ticktin, T., A. N. Whitehead, and H. A. Fraiola. 2006. Traditional gathering of native hula plants in alien-invaded Hawaiian forests: Adaptive practices, impacts on alien invasive species and conservation implications. Environmental Conservation 33:185–194. Uchida, J., and C. Kadooka. 2005. Use of sustainable agricultural practices for taro, Colocasia esculenta in Hawaii. Phytopathology 95:S106. Wilmshurst, J. M., T. L. Hunt, C. P. Lipo, and A. J. Anderson. 2011. High-precision radiocarbon dating shows recent and rapid initial human colonization of East Polynesia. Proceedings of the National Academy of Sciences 108:1815–1820. Zerega, J. C., D. Ragone, and T. J. Motley. 2004. Complex origins of breadfruit (Artocarpus altilis, Moraceae): Implications for human migrations in Oceania. American Journal of Botany 91:760–766. CHAPTER 7. ALIEN INVADERS

Allen, J. A. 1998. Mangroves as alien species: The case of Hawaii. Global Ecology and Biogeography Letters 7:61–71. Ansari, S., and C. C. Daehler. 2010. Life history variation in a temperate plant invader, Verbascum thapsus along a tropical elevational gradient in Hawaii. Biological Invasions 12:4033–4047. Athens, J. S. 2009. Rattus exulans and the catastrophic disappearance of Hawai‘i’s native lowland forest. Biological Invasions 11:1489–1501. Christenhusz, M. J. M., and T. K. Toivonen. 2008. Giants invading the tropics: The oriental vessel fern, Angiopteris evecta (Marattiaceae). Biological Invasions 10:1215–1228. Cox, E. F., and J. A. Allen. 1999. Stand structure and productivity of the introduced Rhizophora mangle in Hawaii. Estuaries 22:276–284. Daehler, C. C. 2006. Invasibility of tropical islands by introduced plants: Partitioning the influence of isolation and propagule pressure. Preslia 78:389–404. ———. 2005. Upper-montane plant invasions in the Hawaiian Islands: Patterns and opportunities. Perspectives in Plant Ecology Evolution and Systematics 7:203–216. Daehler, C. C., and D. A. Carino. 1998. Recent replacement of native pili grass (Heteropogon contortus) by invasive African grasses in the Hawaiian Islands. Pacific Science 52:220–227. D’Antonio, C. M., J. T. Tunison, and R. K. Loh. 2000. Variation in the impact of exotic grasses on native plant composition in relation to fire across an elevation gradient in Hawaii. Austral Ecology 25:507–522. Denslow, J. S. 2003. Weeds in paradise: Thoughts on the invasibility of tropical islands. Annals of the Missouri Botanical Garden 90:119–127. Denslow, J. S., J. C. Space, and P. A. Thomas. 2009. Invasive exotic plants in the tropical Pacific Islands: Patterns of diversity. Biotropica 41:162–170. Funk, J. L. 2005. Hedychium gardnerianum invasion into Hawaiian montane rainforest: Interactions among litter quality,

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Goldman, R. L., L. P. Goldstein, and G. C. Daily. 2008. Assessing the conservation value of a human-dominated island landscape: Plant diversity in Hawaii. Biodiversity and Conservation 17:1765–1781. Goldstein, J. H., G. Caldarone, T. K. Duarte, D. Ennaanay, N. Hannahs, G. Mendoza, S. Polasky, S. Wolny, and G. C. Daily. 2012. Integrating ecosystem-service tradeoffs into land-use decisions. Proceedings of the National Academy of Sciences 109:7565–7570. Gramling, C. 2005. Conservation biology: Hawaii’s coral trees feel the sting of foreign wasps. Science 310:1759–1760. Gutrich, J., D. Donovan, M. Finucane, W. Focht, F. Hitzhusen, S. Manopimoke, D. McCauley, B. Norton, P. Sabatier, J. Salzman, and V. Sasmitawidjaja. 2005. Science in the public process of ecosystem management: Lessons from Hawaii, Southeast Asia, Africa and the US Mainland. Journal of Environmental Management 76:197–209. Hess, S. C., P. C. Banko, G. J. Brenner, and J. D. Jacobi. 1999. Factors related to the recovery of subalpine woodland on Mauna Kea, Hawaii. Biotropica 31:212–219. Hobdy, R. 1993. Lana‘i: A case study: The loss of biodiversity on a small Hawaiian island. Pacific Science 47:201–210. Kearns, C. A., D. W. Inouye, and N. Waser. 1998. Endangered mutualisms: The conservation of plant-pollinator interactions. Annual Review of Ecology and Systematics 29:83–112. Kellner, J. R., G. P. Asner, S. Cordell, J. M. Thaxton, K. M. Kinney, T. Kennedy-Bowdoin, D. E. Knapp, E. J. Questad, and S. Ambagis. 2012. Historical land-cover classification for conservation and management in Hawaiian subalpine drylands. Pacific Science 66:457–466. Kingsford, R. T., J. E. M. Watson, C. J. Lundquist, O. Venter, L. Hughes, E. L. Johnston, J. Atherton, M. Gawel, D. A. Keith, B. G. Mackey, C. Morley, H. P. Possingham, B. Raynor, H. F. Recher, and K. A. Wilson. 2009. Major conservation policy issues for biodiversity in Oceania. Conservation Biology 23:834–840. Kirch, P. V. 2007. Three islands and an archipelago: Reciprocal interactions between humans and island ecosystems in Polynesia. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 98:85–99. Kraus, F., and D. C. Duffy. 2010. A successful model from Hawaii for rapid response to invasive species. Journal for Nature Conservation 18:135–141. Krushelnycky, P. D., L. L. Loope, T. W. Giambelluca, F. Starr, K. Starr, D. R. Drake, A. D. Taylor, and R. H. Robichaux. 2013. Climate-associated population declines reverse recovery and threaten future of an iconic high-elevation plant. Global Change Biology 19:911–922. Loope, L. L. 1995. Climate change and island biological diversity. In P. Vitousek, L. Loope, and H. Adsersen (eds.), Biological Diversity and Ecosystem Function on Islands. Springer-Verlag, New York, pp. 123–134. Loope, L. L., and T. W. Giambelluca. 1998. Vulnerability of island tropical montane cloud forests to climate change, with special reference to East Maui, Hawaii. Climate Change 39:503–517. Loope, L. L., and F. Kraus. 2009. Preventing establishment and spread of invasive species: Current status and needs. In T.

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variation in the erythrina gall wasp (Quadrastichus erythrinae Kim). Pacific Science 64:22–31. Safeeq, M., and A. Fares. 2012. Hydrologic response of a Hawaiian watershed to future climate change scenarios. Hydrological Processes 26:2745–2764. Sakai, A. K., W. L. Wagner, and L. A. Mehrhoff. 2002. Patterns of endangerment in the Hawaiian flora. Systematic Biology 51:276–302. Shiels, A. B., and D. R. Drake. 2011. Are introduced rats (Rattus rattus) both seed predators and dispersers in Hawaii? Biological Invasions 13:883–894. Smith, D. G., E. K. Shiinoki, and E. A. VanderWerf. 2006. Recovery of native species following rat eradication on Mokoli‘i Island, O‘ahu, Hawai‘i. Pacific Science 60:299–303. Thaxton, J. M., T. C. Cole, S. Cordell, R. J. Cabin, D. R. Sandquist, and C. M. Litton. 2010. Native species regeneration following ungulate exclusion and nonnative grass removal in a remnant Hawaiian dry forest. Pacific Science 64:533–544. Ticktin, T. N., A. M. Whitehead, and H. Fraiola. 2006. Traditional gathering of native hula plants in alien-invaded Hawaiian forests: Adaptive practices, impacts on alien invasive species, and conservation implications. Environmental Conservation 33:185–194. Timm, O., and H. F. Diaz. 2009. Synoptic-statistical approach to regional downscaling of IPCC twenty-first century climate projections: Seasonal rainfall over the Hawaiian Islands. Journal of Climate 22:4261–4280. Vitousek, P. 2006. Ecosystem science and human-environment interactions in the Hawaiian archipelago. Journal of Ecology 94:510–521. Wagner, W. L., M. M. Bruegmann, D. R. Herbst, and J. Q. C. Lau. 1999. Hawaiian vascular plants at risk: 1999. Bishop Museum Occasional Papers 60:1–58. Weller, S. G., R. J. Cabin, D. H. Lorence, S. Perlman, K. Wood, T. Flynn, and A. K. Sakai. 2011. Alien plant invasions, introduced ungulates, and alternative states in a mesic forest in Hawaii. Restoration Ecology 19:671–680. Wester, L. 1994. Weed management and the habitat protection of rare species: A case-study of the endemic Hawaiian fern Marsilea villosa. Biological Conservation 68:1–9. Woodcock, D. 2003. To restore the watersheds: Early twentieth-century tree planting in Hawai‘i. Annals of the Association of American Geographers 93:624–635. Yocom, C. F. 1967. Ecology of feral goats in Haleakala National Park, Maui, Hawaii. American Midland Naturalist 77:418–451.

PHOTOGRAPH CREDITS

Note: Unless mentioned below, photographs are by Robert Gustafson.

John K. Obata—4.138, 4.140, 4.141, 4.144, 4.154, 4.166, 4.171, 4.175, 4.185, 4.191 Ocean Conservancy—1.1 Hank Oppenheimer—2.6d, 4.289, 4.619, 5.64, 5.90, 5.114 John Replogle—1.26 Philip W. Rundel—3.18, 8.6 Robert Shallenberger—1.2, 1.6, 1.20, 1.21, 1.22 Forest and Kim Starr—1.15, 2.9, 3.5, 3.10, 3.13, 3.14, 3.25, 3.27, 4.230, 4.386, 4.595, 4.597, 4.635, 4.641, 4.618, 4.622, 4.629, 4.633, 6.6, 6.9, 7.8, 7.16, 8.5 The Nature Conservancy, Hawai‘i—1.14, 1.19, 3.8, 8.9 Philip Thomas—2.2, 2.5a,d Travis Thurston, Wikimedia Commons—1.13 U.S. Fish and Wildlife Service—3.28 U.S. Geological Survey—1.5, 1.23, 1.27, 1.28 Dietrich Varez—6.1 Arthur Whistler—4.223 Ken Wood—4.116 Laurel Woodley—preface, 4.19, 4.350, 4.508, 5.67, 6.2, 7.1

Susan Bainbridge and Bruce G. Baldwin—4.63 Bishop Museum—8.1 Sherwin Carlquist—4.43, 4.126, 4.580 Gerald D. Carr—2.11, 4.58, 4.59, 4.62, 4.67, 4.77, 4.106, 4.146, 4.396, 4.561, 4.621 Sheila Conant—4.3, 4.578 G. Daida—4.612 David Eickhoff—4.394 EPSCOR Hawai‘i—8.7 Mark Fickett, Wikimedia Commons—1.17 Betsy Gagne—1.16, 3.23, 3.24, 8.4 George K. Linney—4.125, 4.608 Arthur C. Medeiros—3.20, 4.78, 4.444, 8.8 Timothy J. Motley—4.165 NOAA–1.3. 1.4

309

Hibiscadelphus hualalaiensis

INDEX

‘a‘ali‘i, 160 Abutilon, 120 eremitopetalum, 120 incanum, 120 menziesii, 120 sandwicense, 120 Acacia, 101 farnesiana, 234 koa, 39, 41, 45, 101, 244, 249 koaia, 101 Acaena, 144 Acaena exigua, 144 Achyranthes, 49–50 splendens, 34, 50 splendens var. rotundata, 50 splendens var. splendens, 50 adaptive radiation, 25 adder’s tongue, 212 Adenophorus, 24, 42, 193, 212 abietinus, 212 epigaeus, 25, 212 hymenophylloides, 26, 212 periens, 213 pinnatifidus, 213 pinnatifidus var. pinnatifidus, 213 pinnatifidus var. rockii, 213 tamariscinus, 26, 213 tamariscinus var. montanus, 213 tamariscinus var. tamariscinus, 213 tenellus, 213 tripinnatifidus, 26, 213 Adenostemma, 57 viscosum, 57 Adiantum, 216 capillus-veneris, 216 a‘e, 156–157, 160 ‘ae, 215 ‘ae lau nui, 215 ‘ae‘ae, 135, 162 African tulip tree, 238 Agavaceae, 170, 172 Agrostis, 183 sandwicensis, 46, 47, 183 ‘ahakea, 145 ‘ahakea lau nui, 145 ‘ahaniu, 178 ‘ahea, 51

‘āheahea, 51 ‘āhihi, 127 ‘āhihi kū ma kua, 127 ‘āhihi lehua, 127 ‘ahina kuahiwi, 201 ‘ahinahina, 50, 58, 59 ‘ahu‘awa, 175 ahupua‘a, 242 a‘ia‘i, 125 ‘aiakanēnē, 145 ‘aiakeakua, 163 ‘aiea, 162–163 Aizoaceae, 49 aka‘awa, 190 ‘ākala, 144 ‘ākalakala, 144 ‘aki, 189 ‘ākia, 164 ‘akiahala, 122 ‘aki‘aki, 189 ‘akoko, 98–101 ‘akole, 202 ‘ākōlea, 165, 200, 221, 222 ‘akū, 83 ‘ākulikuli, 49, 162 ‘ākulikuli kai, 162 ‘akulikuli ‘ōhelo, 162 ‘āla‘a, 161 ‘ala‘ala pū loa, 124 ‘ala‘ala wai nui, 115, 132–133 ‘ala‘alahua, 160 ala‘alai, 221 alahe‘e, 152 Alaka‘i Swamp, 42 Alaka‘i Plateau, 5 alani, 152–155 alani kuahiwi, 152–155 Albizia moluccana, 237 ale, 136 Alectryon, 160 macrococcus, 160 macrococcus var. auwahiensis, 160 macrococcus var. macrococcus, 160 alena, 127 Aleurites moluccana, 35, 39, 228 aloalo, 122 Alocasia, 170 macrorrhizos, 226

Alphitonia, 142 ponderosa, 142 alpine zone, 46–47 Alsinodendron, 87 lychnoides, 88 obovatum, 89 trinerve, 90 viscosum, 90 Alyxia, 53 oliviformis, 53 stellata, 53 Amaranthaceae, 49 Amaranthus, 50 brownii, 50 ‘ama‘u, 197 Amauropelta, 220 globulifera, 221 Anacardiaceae, 52 ‘ānapanapa, 142 ‘ānaunau, 72 Andropogon virginicus, 35, 39, 235–236, 241, 249 Anectochilus, 181 sandvicensis, vi, 181 Angiopteris evecta, 211, 240 ‘ānini, 130 ‘ānounou, 72 Antidesma, 97, 130 platyphyllum, 130 platyphyllum var. hillebrandii, 130 platyphyllum var. platyphyllum, 130 pulvinatum, 130 Antirrhinaceae, 135 ‘ānunu, 93–94 ‘ape, 170, 226 ‘ape‘ape, 112 Apiaceae, 52 Apocynaceae, 53 ‘apu‘u, 198 Aquifoliaceae, 54 Araceae, 170 Arachnoides, 200 insularis, 200 Araliaceae, 55 Arecaceae, 170 Argemone, 129 glauca, 28, 129 glauca var. decipiens, 129

311

glauca var. glauca, 129 Argyroxiphium, 22, 57, 61 calignis, 44, 45, 58 grayanum, 44, 58 kauense, 44, 58 sandwicense, 58 sandwicense subsp. macrocephalum, 22, 26, 46, 58, 59 sandwicense subsp. sandwicense, 46, 59 Artemisia, 59 australis, 59 mauiensis, 59 Artocarpus altilis, 225, 227 Asparagaceae, 172 Aspleniaceae, 193 Asplenium, 24, 42, 193 acuminatum, 193 adiantum-nigrum, 194 caudatum, 194 contiguum, 194 contiguum var. contiguum, 194 dielfalcatum, 194 diellaciniatum, 194 dielpallidum, 194 hobdyi, 195 horridum, 194 kaulfussi, 195 lobulatum, 195 macraei, 195 nidus, 195 normale, 195 peruvianum, 196 peruvianum var. insulare, 196 polyodon, 196 sphenotomum, 196 trichomanes, 196 trichomanes var. densum, 196 Astelia, 45, 173 argyrocoma, 173 menziesiana, 173 Asteliaceae, 173 Asteraceae, 57 Athyriaceae, 196, 223 Athyrium, 222 microphyllum, 222 au, 148, 157 ‘auhuhu, 231 āulu, 128, 160, 161

312

INDEX

aupaka, 168 Australian tree fern, 240 ‘awapuhi, 230 ‘awapuhi-a-Kanaloa, 182, 228 āweoweo, 51 ‘āwikiwiki, 102 ‘āwiwi, 106, 149 Axis axis, 248 axis deer, 35, 248 Axonopus fissifolius, 52 Bacopa, 135, 161 monnieri, 135 balsma pear, 93 banana, 227 banana poka, 239 bastard sandalwood, 161 beach dropseed, 189 beach morning glory, 93 beach pea, 105 beach she-oak, 234 beach vitex, 117 beak-rush, 178 beard grass, 235, 249 beggar’s tick, 59–61 Begoniaceae, 70 bentgrass, 183 Bidens, 22, 26, 37, 59 alba, 59 amplectens, 60 cosmoides, 60 forbesii, 60 forbesii subsp. forbesii, 60 forbesii subsp. kahiliensis, 60 hillebrandiana, 60 hillebrandiana subsp. hillebrandiana, 60 hillebrandiana subsp. polycephala, 60 menziesii, 60 menziesii subsp. filiformis, 60 menziesii subsp. menziesii, 60 molokaiensis, 60 pilosa, 59 sandvicensis, 61 sandvicensis subsp. confusa, 61 sandvicensis subsp. sandwicensis, 61 torta, 61 valida, 61 Big Island. See Hawai‘i birdcatching sedge, 179 bitter yam, 227 black twig borer, 131, 248 black-tailed deer, 248 Blechnaceae, 197 blue gum, 240 blue-eyed grass, 180 Bobea, 145 elatior, 39, 145 sandwicensis, 145 Boehmeria, 165 grandis, 165 nivea, 165 Boerhavia, 127 repens, 34, 127

bogs, 42–45 Bolboschoenus, 174 maritimus, 174 Bombacaceae, 119, 163 Bonamia, 92 menziesii, 92 Boraginaceae, 70 Botrychium subbifoliatum, 211 bottle gourd, 93 bracken fern, 200 Brassicaceae, 71 breadfruit, 225, 227 Brighamia, 20, 72 insignis, ii, 21, 72 rockii, 73 broomsedge, 235, 249 Broussaisia, 113 arguta, 113 Broussonetia papyrifera, 229 Brugiera sexangula, 241 bryophytes, 25 Buddlejaceae, 161 bulrush, 174 Caesalpinia, 101 bondoc, 34, 102 kavaiensis, 101 Calamagrostis, 184 hillebrandii, 184 Callistopteris baldwinii, 205 Calophyllum inophyllum, 229 caltrop, 170 Campanulaceae, 72 Canavalia, 102 galeata, 102 kauaiensis, 102–103 maritima, 102 candlenut, 228 Canthium odoratum, 152 caper family, 87 Capparaceae, 87 Capparis, 87 sandwichiana, 34, 87 Carex, 174 alligata, 174 macloviana, 46 meyenii, 174 montis-eeka, 174 wahuensis, 175 wahuensis subsp. rubiginosa, 175 Carlquista muirii, 22 Caryophyllaceae, 87 Cassytha, 117 filiformis, 118 Casuarina equisetifolia, 35, 234–235 catbrier, 190 catchfly, 91 cattle grazing, 247 Celastraceae, 91 Cenchrus, 184 agrimonioides, 184 agrimonioides var. agrimonioides, 184 agrimonioides var. layysanensis, 184

ciliaris, 35 clandestinus, 35, 39, 236, 241 setaceus, 35, 37, 236, 241 Centaurium sebaeoides, 106 centaury, 106 Chamaesyce, 97 atrococca, 98 celastroides, 98 cluisiifolia, 98 degeneri, 99 deppeana, 99 halemanui, 99 herbstii, 99 multiformis, 100 olowaluana, 100 remyi, 100 rockii, 100 skottsbergii, 100 sparsifolia, 101 Charpentiera, 50 densiflora, 50 elliptica, 50 obovata, 39, 50 tomentosa, 51 tomentosa var. maakuaensis, 51 tomentosa var. tomentosa, 51 Cheirodendron, 44, 55 fauriei, 41, 55 platyphyllum, 41, 55 platyphyllum subsp. kauaiensis, 55 platyphyllum subsp. platyphyllum, 55 trigynum, 41, 55 trigynum subsp. helleri, 55 trigynum subsp. trigynum, 55 Chenopodiaceae, 49, 91 Chenopodium, 51 oahuense, 34, 45, 46, 51 Chinese banyan, 238 Christella, 220 cyantheoides, 220 Christmas berry, 237 Chrysodracon, 223 Chrysopogon, 184 aciculatus, 184 Cibotiaceae, 198 Cibotium, 41, 42, 198 chamissoi, 41, 192, 198 glaucum, 41, 198 menziesii, 41, 198, 199 Citharexylum caudatum, 235 spinosum, 235 Cladium, 175 jamaicense, 175 Claoxylon, 98 sandwicense, 98 Cleomaceae, 87 Cleome, 87 spinosa, 91 Clermontia, 20, 26, 27, 41, 72, 73, 76 arborescens, 21, 73 arborescens subsp. arborescens, 73 arborescens subsp. waihiae, 73 calophylla, 73

clermontioides, 73 clermontioides subsp. clermontioides, 73 clermontioides subsp. rockiana, 73 drepanomorpha, 73 fauriei, 74 grandiflora, 74 grandiflora subsp. grandiflora, 74 grandiflora subsp. maxima, 74 grandiflora subsp. muroi, 74 hawaiiensis, 74 kakeana, 74 kohalae, 74 lindseyana, 74 montis-loa, 75 oblongifolia, 75 oblongifolia subsp. brevipes, 75 oblongifolia subsp. mauienesis, 75 oblongifolia subsp. oblongifolia, 75 pallida, 75 parviflora, 73 peleana, 75 peleana subsp. peleana, 75–76 peleana subsp. singuliflora, 76 persicifolia, 76 tuberculata, 76 waimeae, 76 Clidemia hirta, 238–239, 241, 247 coast sandalwood, 159 coastal shrublands and woodlands, 34 coastal strand and shore cliffs, 33 Coccinea grandis, 235 Coccoloba uvifera, 34 Cocculus, 125 orbiculatus, 125 coconut palm, 34, 229 Cocos nucifera, 34, 229 Colocasia, 170 esculenta, 224, 226 colonization of the Hawaiian Islands, 19 Colubrina, 142 asiatica, 142 oppositifolia, 142, 246 common velvet grass, 241 Coniogramme, 216 pilosa, 216 Convolvulaceae, 92 Coprosma, 154 elliptica, 41, 154 ernodeoides, 45, 47, 145 foliolosa, 146 kauensis, 41, 146 longifolia, 41, 146 menziesii, 146 montana, 45, 146 ochracea, 41, 147 pubens, 147 ternata, 147 waimeae, 41, 147 Cordia, 70 subcordata, 71, 125, 230

INDEX

Cordyline fruticosa, 35, 230 terminalis, 35, 230 Corynocarpus laevigatus, 238 Crepidomanes, 206 draytonianum, 206 minutum, 206 Cressa, 92 truxillensis, 33, 92 Cryptocarya, 118 mannii, 118 Ctenitis, 200 latifrons, 200 Cucumis, 93 Cucurbitaceae, 93 cudweed, 68 Curcuma longa, 230 Cuscuta, 92 Cuscuta sandwichiana, 92 Cuscutaceae, 94 Cyanea, 20, 26, 27, 28, 41, 72, 76 aculeatiflora, 76 acuminata, 77 angustifolia, 77 arborea, 77, 245 asarifolia, 77 calycina, 77 copelandii, 77 copelandii subsp. copelandii, 77 copelandii subsp. haleakalaensis, 77 coriacea, 78 crispa, 78 cylindrocalyx, 79 degeneriana, 78 dolichopoda, 77 dunbariae, 77 elliptica, 77 fissa, 77 floribunda, 79 grimesiana, 79 grimesiana subsp. cylindrocalyx, 79 grimesiana subsp. grimesiana, 79 grimesiana subsp. obatae, 79 hamatiflora, 21, 79 hamatiflora subsp. carlsonii, 79 hamatiflora subsp. hamatiflora, 79 hirtella, 79 horrida, 28 humboldtiana, 79 koolauensis, 80 kunthiana, 80 lanceolata, 80 leptostegia, 80 lobata, 80 lobata subsp. baldwinii, 80 lobata subsp. lobata, 80 longiflora, 81 macrostegia, 81 mceldowneyi, 81 membranacea, 81 pilosa, 81 pilosa subsp. longipedunculata, 81

pilosa subsp. pilosa, 81 pinnatifida, 81 remyi, 82 scabra, 82 solenocalyx, 82 spathulata, 82 st.-johnii, 82 superba, 83 superba subsp. regina, 83 superba subsp. superba, 83 tritomantha, 28, 83 truncata, 82, 83 Cyclosorus, 220 cyantheoides, 220 interruptus, 220 sandwicensis, 221 Cyperaceae, 173 Cyperus, 175 hillebrandii, 175 hillebrandii var. hillebrandii, 175 hypochlorus, 175 hypochlorus var. hypochlorus, 175 javanicus, 175 laevigatus, 176 odoratus, 176 pennatiformis, 176 pennatiformis subsp. bryanii, 176 pennatiformis subsp. pennatiformis, 176 polystachyos, 176 Cyrtandra, 24, 26, 41, 107 calpidicarpa, 107 cordifolia, 107 dentata, 108 garnotiana, 108 grandiflora, 108 hashimotoi, 108 lessoniana, 108 lysiosepala, 108 macrocalyx, 109 oenobarba, 109 paludosa, 109 paludosa var. microcarpa, 109 paludosa var. paludosa, 109 platyphylla, 109 procera, 109 propinqua, 110 sandwicensis, 110 spathulata, 110 waianaeensis, 110 wawrae, 110 Cyrtomium, 200 caryotideum, 201 falcatum, 201 Cystopteridaceae, 223 Cystopteris, 222 douglasii, 222 sandwicensis, 222 damselflies, 20 Daucus pusilla, 52 Degener, Otto, 250 Delissea, 20, 72, 83, 245

argutidentata, 83, 243–244 kauaiensis, 83 rhytidosperma, 84 subcordata var. kauaiensis, 84 undulata subsp. kauaiensis, 84 undulata subsp. undulata, 83 waianaeensis, 84 Dennstaedtiaceae, 199 Deparia, 222 prolifera, 222 Deschampsia, 185 nubigena, 39, 44, 45, 46, 185 Deschampsia grassland, 46 Diamond Head, 7 Dianella, 179, 181, 191 sandwicensis, 191 Dichanthelium, 44, 185 cynodon, 185 isachnoides, 185 Dicksoniaceae, 200 Dicranopteris, 204 linearis, 42, 204 Didiereaceae, 137 Diellia, 24, 193 erecta, 194 falcata, 194 pallida, 195 Digitaria, 185 setigera, 185, 225 dioecy, 26–27 Dioscorea alata, 227 bulbifera, 227 pentaphylla, 227 Diospyros, 95 hillebrandii, 95 sandwicensis, 35, 36, 95 Dipentodontaceae, 95 Diplazium, 222 sandwichianum, 223 Diplopterygium, 205 pinnatum, 205 dispersal, 20 Dissochondrus, 185 biflorus, 185 ditchgrass, 190 dock, 137 dodder, 92 Dodonaea, 160 viscosa, 39, 46, 47, 160, 249 Doodia, 197 Doodia kunthiana, 197 Doryopteris, 216 decipiens, 217 decora, 217 takeuchii, 217 Dracaenaceae, 172 Drosera, 95 anglica, 45, 95 Droseraceae, 95 Drosophila, 20 dry forests, 34–36 Dryopteridaceae, 200 Dryopteris, 24, 42, 201 crinalis, 201 crinalis var. crinalis, 201 crinalis var. podosorus, 201

fusco-atra, 201 fusco-atra var. fusco-atra, 201 fusco-atra var. lamoureuxii, 201 glabra, 201 glabra var. glabra, 201 hawaiiensis, 201 rubiginosa, 202 tetrapinnata, 202 unidentata, 202 unidentata var. palacea, 202 wallichiana, 202 Dubautia, 22, 26, 27, 41, 61 arborea, 61 carrii, 61 ciliolata, 22, 26, 62 ciliolata subsp. ciliolata, 62 ciliolata subsp. glutinosa, 62 herbst-obatae, 62 kalalauensis, 62 knudsenii, 62 knudsenii subsp. knudsenii, 62 imbricata, 45 latifolia, 62 laxa, 45, 63 laxa subsp. hirsuta, 63 linearis, 63 linearis subsp. hillebrandii, 63 linearis subsp. linearis, 63 menziesii, 26, 45, 47, 63 microcephala, 63 paleata, 45, 63 plataginea, 63 plantaginea subsp. humilis, 64 plantaginea subsp. magnifolia, 63, 64 plantaginea subsp. plantaginea, 63, 64 platyphylla, 64 raillardioides, 45, 64 reticulata, 64 scabra, 26, 64 scabra subsp. leiophylla, 64 scabra subsp. scabra, 64 syndetica, 64 waialealae, 45, 65 waianapanapaenis, 64 dwarf iliau, 70 dwarf naupaka, 111 Ebenaceae, 95 ebony, 95 Echium, 27 ehu‘awa, 176 ‘ēkaha, 195, 202–204 ‘ēkaha ‘ākōlea, 214 ‘ēkaha kuahiwi, 195 ‘eke silversword, 58 ‘ekoko, 98–101 El Niño–Southern Oscillation (ENSO), 16 Elaeocarpaceae, 96 Elaeocarpus, 96 bifidus, 96 ēlama, 95 Elaphoglossum, 24, 42, 202 aemulum, 202

313

314

INDEX

alatum, 203 crassifolium, 203 fauriei, 203 paleaceum, 203 pellucidum, 203 wawrae, 204 Eleocharis, 176 obtusa, 177 elephant ears, 226 Embelia, 138 pacifica, 138 ‘emoloa, 186 Emperor Seamounts, 1 ‘ena‘ena, 68 Entada, 102 phaseoloides, 102 Epacridaceae, 96 Eragrostis, 186 atropioides, 46, 186 grandis, 186 variabilis, 34, 186 Erechtites, 57 Ericaceae, 96 Erythrina, 28, 102 sandwicensis, 34, 35, 39, 102–103 Eucalyptus, 125 globulus, 240 Eugenia, 125 reinwardtiana, 125 uniflora, 125 Euphorbia, 26, 27, 28, 98 atrococca, 98 celastroides, 37, 98 celastroides var. amplectens, 98 celastroides var. kaenana, 98 clusiifolia, 42, 98 degeneri, 34, 99 deppeana, 99 haeleeleana, 99 halemanui, 99 herbstii, 99 multiformis, 99 multiformis var. microphylla, 99, 100 multiformis var. multiformis, 100 olowaluana, 100 remyi, 42, 100 remyi var. kauaiensis, 100 rockii, 42, 100 skottsbergii, 100 skottsbergii var. skottsbergii, 100 skottsbergii var. vaccinioides, 100 sparsiflora, 101 Euphorbiaceae, 97 Eurya, 130 sandwicensis, 130 everlasting, 68 Exocarpus, 157 gaudichaudii, 157–158 Fabaceae, 101

Falcataria moluccana, 237 false nettle, 165 fern diversity, 24–25 Festuca, 186 aloha, 186 molokaiensis, 186 Ficus microcarpa, 125, 238 fiddlewood, 235 Fimbristylis, 177 cymosa, 34, 177 cymosa subsp. umbellatocapitata, 177 fire, 35, 39, 37, 39, 234, 235–236, 248–249, 250 fire tree, 237 fireweed, 57 fish poison plant, 231 Flacourtiaceae, 105, 157 Florida prickly blackberry, 239 flowering plant diversity, 20 Flueggea, 97, 130 neowawraea, 131 fountain grass, 236 Fragaria, 144 chiloensis, 144 chiloensis subsp. sandwicensis, 144 French Frigate Shoals, 3 Freycinetia, 182 arborea, 182 fringed orchid, 182 fruitflies, 20 Gahnia, 177 beecheyi, 177 gahniiformis, 178 gall wasp, 102–103 Gardenia, 147 brighamii, 147 Gardner Pinnacles, 3 Gentianaceae, 105 Geraniaceae, 106 Geranium, 106 arboreum, 106 cuneatum, 45, 106 cuneatum subsp. cuneatum, 106 cuneatum subsp. hololeucum, 107 cuneatum subsp. tridens, 106–107 hanaense, 107 hillebrandii, 45, 107 humile, 45 multiflorum, 107 Gesneriaceae, 107 ginger, kāhili, 240 white, 240 yellow, 240 Gleicheniaceae, 204 global change, 252–253 Gnaphalium, 65 sandwicensium, 65, 68 goats, 246 golden beardgrass, 184

Gonocormus minutus, 206 proliferus, 206 Goodeniaceae, 110 goosefoot, 51 gorse, 241 Gossypium, 120 tomentosum, 34, 121 Gouania, 143 hillebrandii, 143 meyenii, 143 vitifolia, 143 Gouldia, 148 gourd, 231 Grammitidaceae, 205, 212, 214 Grammitis, 42, 214, 223 hookeri, 214 tenella, 213 gray knickers, 102 greensword, 58 Greensword Bog, 44 Grevillea robusta, 235 Gunnera, 112 kauaiensis, 42, 112 petaloidea, 42, 112 Gunneraceae, 112 Gynochthodes, 148, 150 trimera, 148 hā, 127 ha‘ā, 130 ha‘āmaile, 130 hā‘awa, 134–135 hāhā, 73–83 hahala, 107–109 haili o Pua, 219 hairgrass, 185 ha‘iwale, 107–110 hala, 183 hala pepe, 172–173 hala ‘uhaloa, 124 Haleakalā, 10, 11, 17, 33 hālena, 162–163 Halophila, 180 hawaiiana, 180 Hamakua Coast, 13 hame, 130 Hanauma Bay, 7 hao, 54 Haplopteris, 217 elongata, 217 Haplostachys, 22, 113, 245 haplostachya, 113 hāpu‘u, 198 hāpu‘u i‘i, 199 hāpu‘u pulu, 198 hau, 95, 230 hau hele, 122 hau hele ‘ula, 123 hau hele wai, 122 hau kuahiwi, 121 Hawai‘i, 12–15 Hawai‘i Natural Heritage Program, 251 Hawaiian bamboo, 225, 231 Hawaiian cotton, 120

Hawaiian Ecosystems at Risk project, 251 Hawaiian Islands Marine National Monument, 4 Hawaiian mints, 20 Hawaiian Sugar Planters’ Association, 249 Hawaiian vetch, 105 hea‘e, 156–157 heau, 157 Hedychium coronarium, 240 acuminata, 148 centranthoides, 149 cookiana, 149 elatior, 149 flavescens, 240 fluviatilis, 149 foggiana, 150 formosa, 150 gardnerianum, 240 hillebrandii, 148 knudsenii, 150 littoralis, 150 schlechtendahliana, 149 terminalis, 41, 148 tryblium, 150 Heliotropium, 71 anomalum, 33, 71 curassavicum, 71 Hemerocallidaceae, 179 Hesperocnide, 165 sandwicensis, 165 Hesperomannia, 23, 27, 65 arbuscula, 65 lydgatei, 65 Heteropogon, 186 contortus, 37, 186 heuhiuhi, 104 hi‘aloa, 124 Hibiscadelphus, 121, 245 distans, 121 giffardianus, 121, 245 hualalaiensis, 121, 245, 310 Hibiscus, 121 arnottianus, 122 arnottianus subsp. arnottianus, 122 arnottianus subsp. immaculatus, 122 arnottianus subsp. punaluuensis, 122 brackenridgei, 122 brackenridgei subsp. brackenridgei, 122 brackenridgei subsp. mokuleianus, 122 brackenridgei subsp. molokaiana, 122 clayi, 122 furcellatus, 122 kokio, 122 kokio subsp. kokio, 122 kokio subsp. saintjohnianus, 123 tiliaceus, 95, 230

INDEX

waimeae, 123 waimeae subsp. hannerae, 123 waimeae subsp. waimeae, 123 hihikolo, 102 Hillebrandia, 20, 70 sandwicensis, 24 Hilo ischaemum, 187 Himalayan raspberry, 239 hinahina, 58, 59, 106–107 hinahina kahahai, 71 hō‘awa, 134–135 hoe a Māui, 203 hohiu, 201 hoi, 227 hoi kuahiwi, 190 hō‘i‘o, 223 hō‘i‘o kula, 221 Holcus lanatus, 46, 241 hōlei, 53 hōlio, 118 holly, 54 holly fern, 201 Holoikauaua, 2 hona, 167 honeycreepers, 20–21 honohono, 113 hōpue, 167 Hosmer, R. S., 249 hōwaiaulu, 65 huahekili, 112 huahekili uka, 111 hu‘ahu‘akō, 137 Hualālai, 12 hue, 125 huehue, 125 huehue haole, 240 huluhulu, 121 hulumoa, 157–158 hunakai, 92 Huperzia, 24, 209 erubescens, 209 phyllantha, 209 serrata, 210 hurricanes, 16, 253 hybridization, 26–27 Hydrangea, 113 Hydrangeaceae, 112 Hydrocharitaceae, 179 Hydrophyllaceae, 70, 113 Hylaeus, 248 Hymenasplenium, 196 unilaterale, 196 Hymenophyllaceae, 205 Hymenophyllum, 206 lanceolatum, 206 obtusum, 206 recurvum, 206 Hypolepis, 199 hawaiiensis, 199 ‘Īao Needle, 11 ‘Īao Valley, 10 ‘ie, 182 ‘ie‘ie, 182 ‘ihi, 137–138

‘ihi lā‘au, 211 ‘ihi mākole, 138 ‘ihi‘ihi, 211 ī‘i, 201 i‘iwi, 21 Ilex, 54 anomala, 41, 55 aquifolium, 54 paraguayensis, 54 ‘iliahi, 159 ‘iliahialo‘e, 159 iliau, 69 ‘īlie‘e, 136 ‘ilihia, 109 ‘ilima, 124 ‘ilima pua kea, 120 insular woodiness, 27–28 Ipomoea, 92 alba, 92 batatas, 226 cairica, 226 horsfallii, 92 imperati, 92 indica, 93 littoralis, 93 pes-caprae, 33, 93 violacea, 92 ipu, 231 Iridaceae, 180 ironwood, 234 Isachne, 187 distichophylla, 187 pallens, 187 Ischaemum, 187 byrone, 187 Isodendrion, 167 hosakae, 168 laurifolium, 168 longifolium, 168 Isoëtaceae, 207 Isoëtes, 207 hawaiiensis, 207 ivy scarlet gourd, 93, 235 ‘iwa, 194, 218 ‘iwa puakea, 218 ‘iwa‘iwa, 194–195, 216, 217 ‘iwa‘iwa lau nui, 220 Jacquemontia, 93 ovalifolia, 93 sandwicensis, 93 Java plum, 127 jewel orchid, 181 Joinvillea, 180 ascendens, 180 ascendens subsp. ascendens, 180–181 Joinvilleaceae, 180 Joseph Rock, 36 Juncaceae, 181 kā ‘ape‘ape, 201 kā ‘i‘iwi, 105 kā palupalu o Kanaloa, 103 Ka‘ala, 6, 16, 42

Kadua, 148 acuminata, 148 affinis, 41, 148 axillaris, 148 centranthoides, 149 cookiana, 149 cordata, 149 cordata subsp. cordata, 149 cordata subsp. remyi, 149 cordata subsp. waimeae, 149 elatior, 149 fluviatilis, 149 foggiana, 149 formosa, 150 knudsenii, 150 littoralis, 34, 150 tryblium, 150 kā‘e‘e, 103 kā‘e‘e‘e, 103 kāhili ginger, 240 Kaho‘olawe, 8–10, 17 kahui, 164 kākalaioa, 102 kākonakona, 188 kākuaohi‘iaka, 93 kala, 129 Kalalau Lookout, 5 kalamālō, 186 kalamoho lau li‘i, 218 Kalapana black sand beach, 14 Kalaupapa Peninsula, 7 kalia, 96 kalo, 170, 220, 226 kaluaha, 173 kaluhā, 174 kāmakahala, 118–119 Kamakou, 7 kāmanamana, 57 kamani, 229 kāmanomano, 184 kamapua‘a, 149 kanakaloa, 124 Kanaloa, 103 kahoolawensis, 103, 245, 246 kanawao, 113 kanawao ke‘oke‘o, 107–110 Kānemiloha‘i, 1 Kānepu‘u, 36 kāpana, 114 kapu system, 225 kapua ‘ilio, 211 karaka, 238 Ka‘ū District, 15, 17 Ka‘ū silversword, 58 Kaua‘i, 5–6, 16 kauila, 142 kauila ‘ānapanapa, 142 kauila kukuku, 142 Ka‘ula, 4 kaulu, 54, 128, 160, 161 kaumahana, 157–158 kauna‘oa, 92 kauna‘oa kahakai, 92 kauna‘oa mālolo, 118 kauna‘oa pehu, 118

Kauō, 3 kauokahiki, 127 Kaupō Gap, 10 kaupo‘o, 93 ka‘upu, 204 kava, 228 kāwa‘u, 55, 103, 156 Kawaihae, 17 Kawaikini, 5 kāwelu, 186 kea, 103 keahi, 161 Keahikauō Bog, 43 Kekaha, 16 Keysserria, 45, 65 maviensis, 65 kī, 230 kiawe, 34, 234 kihe, 215 kikawaiō, 220 kikuyu grass, 236 kīlau, 200, 201 Kīlauea, 14–15, 17 Kīlauea Iki, 15 kilioe, 138 kili‘o‘opu, 176 kiolohia, 176 kīpūkai, 71 klu, 234 ‘kō, 228 koa, 101 koa forest, 38–41 koa haole, 233 koai‘a, 101 koali, 93 koali ‘awa, 93 koali‘ai, 226 koa‘oha, 101 Kohala Mountains, 12, 17 Kohala Volcano, 12 kohekohe, 177 koholāpehu, 62 Kokia, 123 cookei, 123, 245 drynarioides, 123 kauaiensis, 123 koki‘o, 123 koki‘o ke‘oke‘o, 122 koki‘o ‘ula, 122 kokio ‘ula ‘ula, 122 koko, 98–101 Koko Crater, 7 Kōko Seamount, 1 kōkōmālei, 98 kōlea, 140, 141 kōlea lau li‘i, 141 kōlea lau nui, 140 kolekole lehua, 139 koli‘i, 86 kolokolo, 213 kolokolo kahakai, 117 kolokolo kuahiwi, 139 kolokolo mokihana, 153 kolomona, 104 Kona Coast, 16, 17

315

316

INDEX

Kona storms, 16 ko‘oko‘olau, 60, 61 ko‘olau, 60, 61 Ko‘olau Gap, 10 Ko‘olau Mountains, 6, 16 ko‘oloa kea, 120 ko‘oloa ‘ula, 120 kopa, 149 kōpiko, 151–152 kōpiko kea, 151 kōpiko ‘ula, 151 Korthalsella, 158 complanata, 158 latissima, xiii, 158 platycaula, 158 remyana, 158 Koster’s curse, 239 kou, 71, 230 kūau, 195 kuawāwaenohu, 88 kūkaemoa, 153 kūkaenēnē, 145 kukui, 228 kulu‘ī, 51 kūmakanii, 160 kumuniu, 217 kūnānā, 72 kuolohia, 178 kūpala, 94 kūpaoa, 61–65 kupukupu, 208 Kure Atoll, 1, 2, 19 La Pérouse Bay, 10 Labordia, 41, 118 hedyosmifolia, 118–119 hirtella, 118 hosakana, 118 kaalae, 119 tinifolia, 119 tinifolia var. lanaiensis, 119 tinifolia var. tinifolia, 119 tinifolia var. wahiawaensis, 119 venosa, 119 waialealae, 119 waiolani, 119 Lachnagrostis, 187 filiformis, 187 Lagenaria siceraria, 93, 231 Lagenifera, 65 Lagenifera maviensis, 65 Lahaina, 17 Lake Laysan, 3 lama, 95 Lamiaceae, 113 Lāna‘i, 8, 9, 16 Lāna‘i Hale, xiv, 8 Landoltia, 170 lantana, 234 Lantana camara, 35, 234 lapalapa, 55 lau li‘i, 195 laukahi, 202, 208, 218 laukāhi kuahiwi, 135–136 laukahi nunui, 202

Lauraceae, 117 Laysan ‘apapane, 3 Laysan Atoll, 2, 3 Laysan duck, 3 Laysan finch, 2 Laysan millerbird, 3 Laysan rail, 3 Lehua, 4 lehua papa, 126 Lellingeria saffordii, 214, 215 Lemna, 170 Lemnaceae, 181 lepelepe a moa, 219 Lepichinia, 113 hastata, 114 Lepidium, 71 bidentatum, 72 bidentatum var. o-waihiense, 72 serra, 72 Lepisorus, 214 thunbergianus, 214 leponēnē, 145 Leptecophylla, 96 tameiameiae, 38, 39, 45, 47, 96 Lepturus, 187 repens, 34, 188 Leucaena leucocephala, 35, 233, 234, 241 Lichens, 25, 239 Liliaceae, 173, 181 liliwai, 144 Lindsaea, 208 ensifolia, 26, 208 repens, 208 repens var. macraeana, 208 Lindsaeaceae, 207 Liparis, 181 hawaiensis, 182 Lipochaeta, 66 connata, 66 connata subsp. connata, 66 integrifolia, 66 kamolensis, 67 lavarum, 67 lobata, 66 lobata subsp. lobata, 66 micrantha, 67 remyi, 67 rockii, 66, 67 subcordata, 67 succulenta, 66 venosa, 67 Lisianski, 2 Lobelia, 20, 72, 84 dunbariae, 78, 84 dunbariae subsp. paniculata, 84 gaudichaudii, 45 gaudichaudii subsp. koolauensis, 85 gloria-montis, 21, 45, 84 gloria-montis subsp. gloriamontis, 84 gloria-montis subsp. longibracteata, 84 grayana, 21, 85

kauaensis, 45, 48, 85 koolauensis, 85 niihauensis, 85 villosa, 45 yuccoides, 86 lobeliad alliance, 20 Loganiaceae, 118 Lō‘ihi, 14 lola, 193 Lomariopsidaceae, 202, 208 lonomea, 160 loss of herbivore defense, 28 lo‘ulu, 216 loulu, 171–172 loulu hiwa, 172 loulu lelo, 171 lovegrass, 186 Lua Makika, 8 Ludwigia octovalvis, 129, 225 Luzula, 181 hawaiiensis, 46, 181 hawaiiensis var. oahuensis, 181 Lycium, 162 sandwicense, 162 Lycopodiaceae, 209 Lycopodiella, 210 cernua, 210 Lycopodium, 210 Lycopodium venustulum, 210 venustulum var. venustulum, 210 venustulum var. verticale, 210 Lyon, Harold, 237, 238, 249, 250 Lysimachia, 138 glutinosa, 139 hillebrandii, x, 139 kalalauensis, 139 mauritiana, 33, 139 pendens, 139 remyi, 139 venosa, 245 ma‘aloa, 165–166 Machaerina, 177 angustifolia, 177 mariscoides, 178 mariscoides subsp. meyenii, 178 macrofungi, 25 mahiki, 189 māhoe, 160 mai‘a, 227 mai‘a he‘i, 227 mai‘a polapola, 227 maiapilo, 87 maidenhair fern, 216 maile, 53 Mākaha Valley, 6 makaloa, 176 Makapu‘u Beach, 33 mākole, 151 makou, 52 māku‘e, 203 maku‘e lau li‘i, 214 Malvaceae, 119, 163 māmaki, 165

māmane, 104 mamani, 104 mana, 217 mānā, 218 manawahua, 217 mānele, 156–157, 160 manena, 153 manene, 135 mānienie ‘ula, 184 manono, 148 ma‘o, 120, 121 Marattia, 211 Marattia douglasii, 211 Marattiaceae, 210 Mariscus hillebrandii, 175 hypochlorus, 175 javanicus, 176 pennatiformis, 176 Maro Reef, 3 Marsilea, 211 villosa, 34, 211 Marsileaceae, 211 ma‘u, 197 maua, 157 Maui, 10–11, 17 Maui Nui, 7, 19 ma‘uma‘u, 197 Mauna Kea, 12–13, 33 Mauna Loa, 14–15, 33 Maunalei Gulch, 8 mau‘u ‘aki‘aki, 177 mau‘u hō‘ula ‘ili, 180 mau‘u lā‘ili, 180 Mecodium recurvum, 207 mehame, 130 mehamehame, 129, 130 Melanthera, 22, 26, 66 integrifolia, 34, 66 kamolensis, 66 lavarum, 67 micrantha, 67 micrantha subsp. micrantha, 67 remyi, 67 subcordata, 67 venosa, 67 Melastomataceae, 238 Melicope, 24, 152 anisata, 152 barbigera, 153 christophersenii, 41 clusiifolia, 153 cruciata, 41 feddei, 41, 153 haleakalae, 41 hawaiensis, 153 kaalaensis, 153 makahae, 154 oahuensis, 41, 154 orbicularis, 41 ovata, 154 peduncularis, 154 puberula, 154 radiata, 41 rotundifolia, 155 sessilis, 155

INDEX

waialealae, 155 zahlbruckneri, 41 Melinis minutiflora, 39, 236, 249 Menispermaceae, 124 mesic seasonal forest, 38–39 Metrosideros, 42, 44, 126 macropus, 126, 288 polymorpha, 23–24, 36, 37, 38, 39, 46, 47, 126, 244, 249 polymorpha var. dieteri, 24, 126 polymorpha var. glaberrima, 23, 41, 126 polymorpha var. incana, 23, 39, 126 polymorpha var. macrophylla, 23, 126 polymorpha var. newellii, 24, 126 polymorpha var. polymorpha, 41, 126 polymorpha var. pseudorugosa, 23, 44, 126 polymorpha var. pumila, 23, 44, 126 rugosa, 126 tremuloides, 127 waialealae, 44, 127 waialealae var. fauriei, 127 waialealae var. waialealae, 127 Mezoneuron, 103 kavaiense, 101, 103 Miconia calvescens, 238–239, 241 Microlepia, 199 strigosa, 199 strigosa var. mauiensis, 199 strigosa var. strigosa, 199 Microsorum, 214 spectrum, 214 spectrum var. pentadactylum, 214 spectrum var. spectrum, 214 Mid Camp Bog, 44 Midway Atoll, 1 mikinalo, 95 milo, 124 moa, 109, 215–216 moa nahele, 215 moa nalo, 28 mohihihi, 105 mokehana, 152 mokihana kūkae moa, 153 Mokulē‘ia, 36 Mokumanamana, 4 Mokupāpapa, 3 molasses grass, 236, 249 Moloka‘i, 8–9, 17 Moluccan albizia, 237 Monocotyledons, 170 montane wet forest, 40–42 Monterey pine, 240 Montiaceae, 137 Mo‘omoni, 7, 34 Moraceae, 125 Morella faya, 39, 237, 241 Morelotia, 178 gahniiformis, 178

Morinda citrifolia, 35, 231 trimera, 148, 150 Mormodica, 93 morning glory, 92, 226 Mount ‘Eke, 11, 42, 43 Mount Wai‘ale‘ale, 5, 16 mountain apple, 127, 228 mountain pili, 189 Mucuna, 103 benedettii, 103 gigantea, 103 mule deer, 248 mule’s-foot fern, 240 Munroidendron, 55 racemosum, 56 Musa troglodytarum, 227 x paradisiaca, 227 Myoporaceae, 125, 161 Myoporum, 161 sandwicense, 34, 37, 39, 45, 46, 161 Myrica faya, 39, 237 Myrsine, 41, 140 alyxifolia, 140 helleri, 140 kauaiensis, 39 lanaiensis, 140 lessertiana, 41, 140 linearifolia, 141 punctata, 141 sandwicensis, 141 wawraea, 141 Myrtaceae, 125 na‘ena‘e, 61–65 na‘ena‘e pua kea, 63 na‘ena‘e pua melemele, 63 na‘ena‘e ‘ula, 64 naio, 161 Nalukakala, 3 Nama, 71 sandwicensis, 71 Nānākuli Valley, 6 nani Wai‘ale‘ale, 144, 169 nānū, 147 Nāpali Coast, 5 Nasturtium sarmentosum, 72 nā‘ū, 147 naupaka, 112 naupaka kahakai, 112 naupaka kai, 112 naupaka kuahiwi, 111–112 Necker Island, 4 nehe, 66 neleau, 52 nena, 71, 127 neneleau, 52 Nephrolepidaceae, 211, 208 Nephrolepis, 208 cordifolia, 208 exaltata, 209 exaltata subsp. hawaiiensis, 209 Neraudia, 35, 165 kauaiensis, 165 melastomifolia, 166

Nertera, 150 granadensis, 151 Nestegis, 128 sandwicensis, 36, 37, 39, 129 Neva Shoals, 2 New Zealand laurel, 238 ni‘ani‘au, 209 nightshade, 163 Nīhoa, 3 Nīhoa finch, 4 Nīhoa millerbird, 4 Ni‘ihau, 4, 16 nīoi, 125 niu, 229 nohoanu, 106–107 nohu, 170 nohunohu, 170 noni, 231 noni kuahiwi, 148 Northwestern Hawaiian Islands, 1, 19 Nostoc, 112 Nothocestrum, 162 breviflorum, 162, 246 latifolium, 162 longifolium, 162 peltatum, 163 Nothoperanema, 204 Nototrichium, 51 humile, 51 sandwicense, 37, 51 nuku, 105 nuku ‘i‘iwi, 105 nutrush, 179 Nyctaginaceae, 127 o‘a, 142 O‘ahu, 6–7, 16 ōali, 218 ‘oāli‘i, 196 ‘oāli‘i makali‘i, 219 Ochrosia, 53 elliptica, 54 haleakalae, 53 octopus tree, 238 Odontosoria, 208 ‘oha, 83–84 ‘ōhā wai, 73–76 ‘ōhā wai nui, 73 ‘ohai, 104 ‘ohe, 56, 180, 187, 225, 231 ‘ohe kiko‘olā, 57 ‘ohe makai, 56 ‘ohe mauka, 56 ‘ohe naupaka, 111 ‘ohe ‘ohe, 56 ‘ōhe‘e, 152 ‘ōhelo, 97 ‘ōhelo kai, 162 ‘ōhelo kau lā‘au, 97 ‘ōhelo papa, 144 ‘ohe‘ohe, 217 ‘ōhi‘a, 38–41, 126 ‘ōhi‘a ‘āhihi, 127 ‘o‘hia ‘ai, 228

‘ōhi‘a hā, 127 ‘ōhi‘a kū, 206 ‘ōhi‘a lehua, 126 ‘ōkolemakili, 105 ōkupukupu, 197 ōkupukupu lau‘i‘i, 197 ‘olapa, 55 Oleaceae, 128 ‘ōlena, 147, 230 ‘ōliko, 140 olive family, 128 ‘oloa, 165–166 olomea, 95 olonā, 167 olopua, 129 olua, 199 ‘ōlulu, 72 Onagraceae, 129 ‘opeha, 202 Ophioderma, 211 pendulum, 212 Ophioglossaceae, 211 Ophioglossum, 211 pendulum, 212 polyphyllum, 212 ‘ōpiko, 151 ōpuhe, 167 Opuntia ficus-indica, 234 Orchidaceae, 181 Oreobolus, 44, 178 furcatus, 178 Oreogrammitis, 223 oriental mangrove, 241 Orobanchaceae, 161 orographic effects, 16–17 Osteomeles, 144 anthyllidifolia, 34, 37, 144, 249 Ovis aries orientalis, 248 musimon, 248 Oxalidaceae, 129 Oxalis, 129 corniculata, 129, 225 Oxyspora paniculata, 239 Pa‘aiki Valley, 34 Pacific Decadal Oscillation (PDO), 16 pai, 212 pā‘ihi, 127 pa‘iniu, 173 pakaha, 114 pākahakaha, 214 pala, 211 pala‘ā, 208 palai, 199 palai hinahina, 206 palai huna, 212 palai lā‘au, 213 palai lau li‘i, 206 palalalai ‘aumakua, 201 palapala‘ā, 208 palapalai, 199 palapalai a Kamapua‘a, 221 pāmakani, 122 pāmakani māhū, 131

317

318

INDEX

pāmoho, 196, 197, 209 pānaunau, 86 Pandanaceae, 182 Pandanus, 183 tectorius, 35, 125, 183 panic grass, 188 Panicum, 188 beecheyi, 188 tenuifolium, 46 torridum, 188 pānini, 234 pāpa‘ahekili, 111 Papa‘āpoho, 2 Papahānaumokuākea, 4 pāpala, 50, 128 pāpala kēpau, 128 papa‘oi, 204 Papaveraceae, 129 paper mulberry, 229 Passiflora mollissima, 239 suberosa, 240 tarminiana, 63, 39–240, 247 pā‘ū o Pala‘e, 208 pā‘ūohi‘iaka, 93 pāwale, 137 pe‘ahi, 214 Pearl and Hermes Atoll, 2 Pelea anisata, 153 barbigera, 153 clusiifolia, 153 feddei, 153 hawaiensis, 153 kaalaensis, 153 makahae, 154 oahuensis, 154 ovata, 154 parvifolia, 155 peduncularis, 154 puberula, 154 rotundifolia, 155 waialealae, 155 Pellaea, 217 ternifolia, 218 Penguin Bank, 7 Pennisetum, 35, 184 clandestinum, 236 setaceus, 236 Pentaphylacaceae, 129, 163 Pēpē‘ōpae Bog, 8, 42, 43 Peperomia, 132 cookiana, 132 globulanthera, 132 hesperomannii, 132 hypoleuca, 132 macraeana, 132 membranacea, 133 obovatilimba, 133 remyi, 133 subpetiolata, 133 tetraphylla, 133 peppergrass, 72 Perispermum, 92 Perrottetia, 91, 95 sandwicensis, 95 persimmon, 95

Peucedanum, 52 sandwicense, 52 Phrymaceae, 161 Phyllanthaceae, 97, 130 Phyllanthus, 97, 131 distichus, 131 Phyllostegia, 22 ambigua, 114 floribunda, 114 grandiflora, 18, 21, 114 hirsuta, 114 lantanoides, 115 variabilis, 34 vestita, 115 Phytolacca, 131 octandra, 131 sandwicensis, 131 Phytolaccaceae, 131 pi‘a, 227 pia, 228 pig, European, 247 Polynesian, 247 Pihemanu, 1 pi‘ipi lau manamana, 195 pi‘ipi‘i, 184 Pilea, 166 peploides, 166 pili, 186 pili grass, 37, 186 pili uka, 189 pilo, 87, 145–149 pilo kea, 156 pilo kea lau li‘i, 156 Pinchot, Gifford, 249 Pinus radiata, 240 pioi, 190 Piper betel, 132 methystichum, 132, 238 methysticum, 228 nigrum, 131 Piperaceae, 131 pipi, 216 pīpīwai, 177 Pipturus, 166 albidus, 39, 166 kauaiensis, 166 Pisonia, 128 brunoniana, 128 grandis, 128 sandwichensis, 128 umbellifera, 39, 128 pitcher sage, 114 Pittosporaceae, 133 Pittosporum, 134 confertiflorum, 134 gayanum, 134 glabrum, 134 hosmeri, 134 kauaiense, 134 terminalioides, 135 Planchonella, 161 sandwicensis, 161 Plantaginaceae, 135, 161 Plantago, 27, 135 hawaiensis, 135

pachyphylla, 135–136 princeps, 136 princeps var. anomala, 136 princeps var. laxifolia, 136 princeps var. longibracteata, 136 princeps var. princeps, 136 psyllium, 135 plantain, 135 Platanthera, 182 holochila, 182 Platydesma, 24, 155 cornuta, 155 cornuta var. cornuta, 155 cornuta var. decurrens, 155 rostrata, 156 spathulata, 156 Plectranthus, 115 parviflorus, 115 Pleomele, 172 aurea, 172 hawaiiensis, 173 Pluchea, 57 Plumbaginaceae, 136 Plumbago, 136 zeylanica, 34, 136 Pneumatopteris sandwicensis, 221 Poa, 188 sandwicensis, 46, 188 siphonoglossa, 189 po‘a‘aha, wauke, 229 Poaceae, 183 po‘e, 138 Pōhakuloa, 37 pohe hiwa, 169 pohinahina, 117 pōhinahina, 71 pōhole, 223 pōhuehue, 93 pōkalakala, 129 pokeberry, 131 pokeweed, 131 pollination, 20, 27 pololei, 212 Polycias, 55 hawaiensis, 56 kavaiensis, 56 oahuensis, 56 racemosa, 56 sandwicensis, 35, 37, 56 waialeale, 57 waimeae, 57 Polygonaceae, 136 Polynesian arrowroot, 228 Polynesian colonization, 101, 183, 225, 246 Polynesians, 19, 92, 233, 245, 248 Polypodiaceae, 212 Polypodium, 215 pellucidum, 215 pellucidum var. pellucidum, 215 pellucidum var. volcanicum, 215 polypody fern, 215

Polystichum, 2, 201, 204 bonseyi, 204 hillebrandii, 204 po‘olā, 98 po‘olā nui, 60 pōpolo, 131, 163 poppy family, 129 portia tree, 124 Portulaca, 137 lutea, 137 molokiniensis, 138 oleracea, 137 sclerocarpa, 138 villosa, 138 Portulacaceae, 137 Potamogeton, 189 nodosus, 190 Potamogetonaceae, 189 Pouteria sandwicensis, 161 prickly poppy, 129 Primulaceae, 125, 138 Pritchardia, 3, 39, 171, 246 arecina, 171 beccariana, 171 hillebrandii, 171 lowreyana, 171 martii, 172 minor, 172 remota, 4, 172 Prosopis pallida, 34, 35, 234 Pseudognaphalium, 65, 67 sandwicensium var. molokaiense, 68 Pseudophegopteris, 221 keraudreniana, 221 Psidium cattleianum, 237, 241, 247 Psidium guajava, 237, 238 Psilotaceae, 215 Psilotum, 215 complanatum, 215 nudum, 216 Psychotria, 26, 151 hawaiiensis, 38 hawaiiensis var. hawaiiensis, 151 hawaiiensis var. hillebrandii, 151 hexandra, 151 hexandra var. hexandra, 151 hexandra var. oahuensis, 151 hobdyi, 151 kaduana, 39, 151 mariniana, 38, 39, 151 mauiensis, 152 Psydrax, 152 odorata, 36, 37, 39, 152 Pteralyxia, 54 kauaiensis, 54 macrocarpa, 54 Pteridaceae, 216 Pteridium, 199 aquilinum, 45, 46, 200 aquilinum, subsp. decompositum, 200

INDEX

Pteris, 218 cretica, 218 excelsa, 218 hillebrandii, 218 irregularis, 218 pū aupaka, 73 pū hala, 183 pua, 129 pua ‘ala, 73 pua kala, 82, 129 pua maka nui, 70 pua‘a olomea, 95 pua‘akuhinia, 173 puakauhi, 102 puaokama, 94 puapua moa, 212 pu‘e, 85 Pūhāhonu, 3 pūhili, 105 puhuehue, 93 pūkiawe, 96 pūnana manu, 196 Punchbowl Crater, 7 puo pilo, 87 pu‘uka, 176 pu‘uko‘a, 178 Pu‘ukukui, 42 Pycreus polystachyos, 176 Ranunculaceae, 141 Ranunculus, 142 hawaiensis, 142 rarity, 244 rat, black, 248 Norway, 248 Polynesian, 248 Rattus exulans, 248 norvegicus, 248 rattus, 248 Rauvolfia, 54 sandwicensis, 54 red mangrove, 241 Remya, 68 kauaiensis, 68 mauiensis, 68 montgomeryi, 68 Reynoldsia, 55 sandwicensis, 35, 56 Rhacomitrium lanuginosum, 44 Rhamnaceae, 142 Rhizophora mangle, 241 Rhus, 52 sandwicensis, 52 Rhynchospora, 178 chinensis subsp. spiciformis, 44 rugosa var. lavarum, 44, 178–179 sclerioides, 179 Rivina humilis, 131 Rock, Joseph, 36, 41, 237, 244, 249 Rollandia, 76 angustifolia, 80 crispa, 78 humboldtiana, 79 lanceolata, 80

lanceolata subsp. calycina, 77 longiflora, 81 st.-johnii, 83 Rorippa, 72 sarmentosa, 72 Rosaceae, 143 Rubiaceae, 145 Rubus, 144 argutus, 239, 241 ellipticus, 239 hawaiensis, 28, 144, 239 macraei, 144 Rumex, 137 albescens, 137 giganteus, 137 skottsbergii, 137 Ruppia, 190 maritima, 190 Ruppiaceae, 190 Ruscaceae, 172 Rutaceae, 152 Saccharum officinarum, 228 Sadleria, 24, 193, 197 cyatheoides, 197 pallida, 41, 197 souleyetiana, 197 squarrosa, 198 unisora, 25, 198 Salicaceae, 105, 157 sandalwood, 158, 246 sandbur, 184 Sanicula, 52 purpurea, 52 sandwicensis, 53 Santalaceae, 157 Santalum, 24 ellipticum, 3, 34, 158, 159 freycinetianum, 39, 158, 159 freycinetianum var. lanaiense, 159 freycinetianum var. pyrularium, 159 haleakalae, 159 haleakalae var. haleakalae, 45, 159 haleakalae var. lanaiense, 159 involutum, 158 paniculatum, 158, 159 paniculatum var. paniculatum, 159 paniculatum var. pilgeri, 159 pyrularium, 159 Sapindaceae, 159 Sapindus, 160 oahuensis, 36, 39, 160 saponaria, 39, 160 Sapotaceae, 161 saw-grass, 175 Scaevola, 24, 110 chamissoniana, 39, 111 coriacea, 34, 111 gaudichaudiana, 39, 111 gaudichaudii, 111 glabra, 111

kilaueae, 111 mollis, 112 procera, 112 sericea, 113 taccada, 33, 112 Sceptridium subbifoliatum, 211 Schefflera actinophylla, 238 Schenkia, 105 sebaeoides, 106 Schiedea, 24, 26, 27, 87, 245 adamantis, 87 globosa, 87 hookeri, 88 kaalae, 88 kealiae, 88 ligustrina, 88 lychnoides, 88 lydgatei, 88 mannii, 89 membranacea, 89 menziesii, 89 nuttallii, 89 obovata, 89 salicaria, 90 spergulina, 90 trinervis, 90 verticillata, 90 viscosa, 90 Schinus terebinthifolius, 35, 237, 241 Schizachyrium condensatum, 35, 39, 235–236, 241, 249 Schizaea, 219 robusta, 219 Schizaeaceae, 219 Schizostachyum glaucifolium, 225, 231 Schoenoplectiella, 179 juncoides, 179 Schoenoplectus, 179 juncoides, 179 lacustris, 179 tabernaemontani, 179 Scirpus, 179 Scleria, 179 testacea, 179 screw pine, 183 Scrophulariaceae, 125, 135, 161 sea bean, 103 sea purslane, 49 seagrass, 180 seaside heliotrope, 71 Selaginella, 219 arbuscula, 219 deflexa, 219 Selaginellaceae, 219 Senegalia, 101 Senna, 104 alat, 104 gaudichaudii, 104 Sesbania, 104 tomentosa, 34, 104 Sesuvium, 49 portulacastrum, 33, 49 Setaria, 185

sexual dimorphism, 26–27 Sicyos, 93 anunu, 93 herbstii, 94 hispidus, 94 lasiocephalus, 94 maximowiczii, 94 pachycarpus, 94 waimanaloensis, 94 Sida, 124 fallax, 34, 124 Sideroxylon , 161 polynesicum, 161 Silene, 90 alexandri, 90, 91 cryptopetala, 90, 91 degeneri, 90, 91 hawaiiensis, 90, 91 lanceolata, 90, 91 perlmanii, 90, 91, 245 struthioloides, 47, 90, 91 silk oak, 235 silversword, 58 Silversword Reserve, 47 Sisyrinchium, 180 acre, 180 Smilacaceae, 190 Smilax, 190 melastomifolia, 190 snakeroot, 52 soapberry, 160 Solanaceae, 162 Solanum, 163 nelsonii, 163 sandwicense, 163 Sophonia rufofascia, 248 Sophora, 104 chrysophylla, 37, 39, 45, 46, 47, 104 Sophora/Myoporum woodland, 46–47 sorrel, 137 South Point, 17 Spanish needles, 59 Spathodea campanulata, 232, 238 Spermolepis, 53 hawaiiensis, viii, 53 Sphaerocionium lanceolatum, 206 obtusum, 206 Sphaeropteris cooperi, 240, 241 Sphagneticola, 57 Sphagnum, 44 Sphenomeris, 208 chinensis, 26, 208 Spirodela, 170 polyrhiza, 170 Sporobolus, 189 virginicus, 33, 189 spurflower, 115 spurge, 98–101 Stachys, 113 Stenogrammitis, 215–216 saffordii, 215 Stenogyne, 22, 26, 41, 115 angustifolia, 115

319

320

INDEX

calaminthoides, 115 kaalae, 116 kaalae subsp. kaalae, 116 kamehamehae, 116 kanehoana, 116 kealiae, 116 microphylla, 116 purpurea, 117, 254 purpurea var. kealiae, 116 rugosa, 117 sessilis, 117 Sterculiaceae, 119, 163 Sticherus, 205 owhyhensis, 206 Stilbaceae, 161 strangler fig, 238 Straussia, 151 strawberry, 144 strawberry guava, 237 Streblus, 125 pendulinus, 125 Strongylodon, 105 nova-guianensis, 105 ruber, 105 Styphelia tameiameiae, 96 subalpine shrubland, 32, 45–46 sugarcane, 228 sumac, 52 sundew, 95 sweet hoary abutilon, 120 sweet potato, 226 Syzygium, 127 jambos, 127 malaccense, 127, 228 sandwicense, 127 Tacca leontopetaloides, 35, 228 Talinaceae, 137 Talipariti tiliaceum, 230 Tanager Peak, 4 tanglehead, 186 Tantalus, 7, 16 taro, 170, 226 teasel gourd, 93 Tectaria, 220 gaudichaudii, 220 Tectariaceae, 219 Tephrosia pupurea, 231 Terminalia catappa, 34 Ternstroemiaceae, 130, 163 Tetramalopium, 22–23, 26, 68 arenarium, 68 arenarium subsp. arenarium, 68 arenarium subsp. laxum, 69 consanguineum, 68 consanguineum subsp. consanguinium, 69 consanguineum subsp. leptophyllum, 69 humile, 47, 69 humile subsp. haleakalae, 69 humile subsp. humile, 69 lepidotum, 69 lepidotum subsp. arbusculum, 69

lepidotum subsp. lepidotum, 69 rockii, 69 rockii var. rockii, 69 sylvae, 69 Tetraplasandra, 55 hawaiensis, 56 kavaiensis, 56 oahuensis, 56 waialealae, 57 waimeae, 57 Theaceae, 130, 163 Thelypteridaceae, 220 Thelypteris, 221 globulifera, 221 Thespesia, 124 populnea, 35, 124. 225 Thymeleaceae, 163 ti, 230 Tibouchina herbacea, 239 urvilleana, 239 Tiliaceae, 119, 163 Timonius, 145 Torulinium odoratum, 176 Touchardia, 166 latifolia, 167 Tournefortia argentea, 34, 70 trade winds, 16 Trematolobelia, 20, 72, 86 kaalae, 86 kauaiensis, 86 macrostachys, 86 macrostachys var. kaalae, 21, 86 singularis, 86 Tribulus, 170 cistoides, 170 Trichomanes Trisetum, 189 glomeratum, 45, 46, 47, 189 turmeric, 230 tussock grassland, 46 twayblade, 182 twisted beardgrass, 186 two-spotted leafhopper, 248 uahiapele, 153 ‘uala, 226 ‘uhaloa, 124 uhauhakō, 137 uhi, 227 uhiuhi, 103 ‘uki, 175, 177–178, 190 ‘uki‘uki, 190 ‘ūlei, 144 Ulex europaeus, 241 ‘ulu, 225 uluhe, 204 uluhe lau nui, 205 Ulupalakua dry forest, 36 ulupua, 129 umbrella sedge, 175, 176 Uncinia, 179 brevicaulis, 46 uncinata, 179 unuhe, 204 Urera, 167

glabra, 167 kaalae, 167 Urticaceae, 165 Vaccinium, 44, 96 berberidifolium, 97 calycinum, 97 dentatum, 97 reticulatum, 39, 45, 46, 97 Vachellia, 101 farnesiana, 234 Vandenboschia, 207 draytoniana, 206 cyrtotheca, 207 davallioides, 207 velvet tree, 239 Verbascum thapsus, 241 Verbenaceae, 167 Veronicaceae, 135 Vespula pensylvanica, 248 Vicia, 105 menziesii, 105 Vigna, 105 adenantha, 105 marina, 33, 105 Viola, 45, 168 chamissoniana, 168 chamissoniana subsp. chamissoniana, 169 chamissoniana subsp. robusta, 169 chamissoniana subsp. tracheliifolia, 169 helenae, 169 kauaensis, 169 kauaensis var. kauaensis, 169 kauaensis var. wahiawaensis, 169 maviensis, 169 wailenalenae, 170 Violaceae, 167 Viscaceae, 157, 170 Vitex, 117 Vitex rotundifolia, 33, 117 Vittariaceae, 223 Wahiawa Bog, 42, 44 wahine noho mauna, 213 Wai‘ale‘ale, 5, 42 Wai‘anae Mountains, 6 Wailau, 7 waimakanui, 218, 221 Waimea Canyon, 5 walahe‘e, 152 Waltheria, 124, 163 indica, 124 wānini, 130 water clover fern, 211 water hyssop, 135 watershed partnerships, 250 wāwae‘iole, 209 Wedelia, 26, 57 widgeon grass, 190 Wikstroemia, 26, 28, 37, 163 bicornuta, 164

furcata, 164 oahuensis, 39, 164 oahuensis var. oahuensis, 164 oahuensis var. palustris, 164 phillyreifolia, 164 pulcherrima, 164 sandwicensis, 46 uva-ursi, 164 uva-ursi var. kauaiensis, 165 uva-ursi var. uva-ursi, 164, 165 wild ginger, 230 wild yam, 227 wiliwili, 102 Wilkesia, 22, 69 gymnoxiphium, 22, 70 hobdyi, 70 wood rush, 181 Woodsiaceae, 196, 221, 223 wooly mullein, 241 Xanthorrhoeaceae, 190 xLindsaeosoria flynii, 208 Xylosandrus compactus, 131, 248 Xylosma, 105, 157 hawaiiense, 37, 157 yam, 227 yellow-faced bees, 248 yellowjacket, 248 Zanthoxylum, 156 dipetalum, 156 dipetalum var. dipetalum, 156 dipetalum var. tomentosum, 156 hawaiiense, 156 kauaense, 157 Zingiber zerumbet, 230 Zygophyllaceae, 170

ABOUT THE AUTHORS

ROBERT J. GUSTAFSON, now retired,

is a former staff botanist at the Los Angeles County Museum of Natural History. He is the coauthor with S. H. Sohmer of Plants and Flowers of the Hawaiian Islands, published by the University of Hawai‘i Press in 1987, and now in at least its 5th edition. He has photographed plants throughout the Hawaiian Islands for more than 30 years. He is the coauthor with Philip Rundel of Introduction to the Plant Life of Southern California: Coast to Foothills (University of California Press, 2005).

DERRAL R. HERBST, now retired,

worked for the U.S. Fish and Wildlife Service and the U.S. Army Corps of Engineers in Hawai‘i for more than 25 years. He is the coauthor of Manual of the Flowering Plants of Hawai‘i, published by the University of Hawai‘i Press, 1990, with Warren L. Wagner and S. H. Sohmer, and A Tropical Garden Flora: Plants Cultivated in the Hawaiian Islands and Other Tropical Places (Bishop Museum, 2005) with George W. Staples.

PHILIP W. RUNDEL is Distinguished

Professor of Biology in the Department of Ecology and Evolutionary Biology at the University of California, Los Angeles, and Director of the Mildred E. Mathias Botanical Garden at UCLA. He is the coauthor or editor of seven previous books and more than 300 research articles (Cambridge University Press, Springer Verlag, Chapman and Hall, University of California Press).

production notes for

Gustafson / HAWAIIAN PLANT LIFE Book design and composition by Mardee Melton, in 9-point Minion Pro, with display type in ITC Avant Garde Gothic. Printing and binding by Regent Publishing Services. Printed on 128 gsm matte art