Trees of Panama and Costa Rica [Course Book ed.] 9781400836178

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Trees of Panama and Costa Rica

Pr in c et o n Fie l d Gu id es

r ooted in field experience and scientific study, Princeton’s guides to animals and plants are the authority for professional scientists and amateur naturalists alike. Princeton Field Guides present this information in a compact format carefully designed for easy use in the field. t he guides illustrate every species in color and provide detailed information on identification, distribution, and biology. Albatrosses, Petrels, and Shearwaters of the World, by d erek o nley and Paul scofield Birds of Australia, eighth edition, by Ken simpson and n icolas d ay Birds of Borneo: Brunei, Sabah, Sarawak, and Kalimantan, by susan Myers Birds of Chile, by Alvaro Jaramillo Birds of the Dominican Republic and Haiti, by steven l atta, c hristopher r immer, Allan Keith, James Wiley, Herbert r affaele, Kent McFarland, and eladio Fernandez Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, and Burundi, by t erry stevenson and John Fanshawe Birds of Europe, by Killian Mullarney, l ars svensson, d an Zetterström, and Peter J. Grant Birds of Europe, second edition, by l ars svensson, d an Zetterström, and Killian Mullarney Birds of India, Pakistan, Nepal, Bangladesh, Bhutan, Sri Lanka, and the Maldives, by r ichard Grimmett, c arol inskipp, and t im inskipp Birds of Kenya and Northern Tanzania: Field Guide Edition, by d ale A. Zimmerman, d onald A. t urner, and d avid J. Pearson Birds of the Middle East, by r . F. Porter, s . c hristensen, and P. schiermacker-Hansen Birds of Nepal, by r ichard Grimmett, c arol inskipp, and t im inskipp Birds of Northern India, by r ichard Grimmett and t im inskipp Birds of Peru, by t homas s . schulenberg, d ouglas F. stotz, d aniel F. l ane, John P. o ’n eill, and t heodore A. Parker iii Birds of the Seychelles, by Adrian s kerrett and ian Bullock Birds of Southeast Asia, by c raig r obson Birds of Southern Africa, by ian sinclair, Phil Hockey, and Warwick t arboton

Birds of Thailand, by c raig r obson Birds of the West Indies, by Herbert r affaele, James Wiley, o rlando Garrido, Allan Keith, and Janis r affaele Birds of Western Africa, by n ik Borrow and r on d emey Caterpillars of Eastern North America: A Guide to Identification and Natural History, by d avid l . Wagner Coral Reef Fishes, by ewald l ieske and r obert Meyers Dragonflies and Damselflies of the West, by d ennis Paulson Mammals of Europe, by d avid W. Macdonald and Priscilla Barrett Mammals of North America, by r oland W. Kays and d on e. Wilson Minerals of the World, by o le Johnsen Nests, Eggs, and Nestlings of North American Birds, second edition, by Paul J. Baicich and c olin J. o . Harrison Palms of Southern Asia, by Andrew Henderson Parrots of the World, by Joseph M. Forshaw The Princeton Field Guide to Dinosaurs, by Gregory s . Paul Raptors of the World, by James Ferguson-l ees and d avid A. c hristie Seeds of Amazonian Plants, by Fernando c ornejo and John Janovec Sharks of the World, by l eonard c ompagno, Marc d ando, and s arah Fowler Stars and Planets: The Most Complete Guide to the Stars, Planets, Galaxies, and the Solar System, Fully r evised and expanded edition, by ian r idpath and Wil t irion Trees of Panama and Costa Rica, by r ichard c ondit, r olando Pérez, and n efertaris d aguerre Whales, Dolphins, and Other Marine Mammals of the World, by Hadoram s hirihai and Brett Jarrett

r ic HAr d c o n d it r ol An do Pér eZ n e Fe rt Ar i s d AG u e rr e

Trees of Panama and Costa Rica

PRINCETON UNI VERSIT Y PRESS

PRINCETON AND OXFORD

Copyright © 2011 by Princeton University Press Published by Princeton University Press, 41 William Street, Princeton, New Jersey 08540 In the United Kingdom: Princeton University Press, 6 Oxford Street, Woodstock, Oxfordshire OX20 1TW nathist.princeton.edu All Rights Reserved Library of Congress Cataloging-in-Publication Data Condit, Richard, 1956– Trees of Panama and Costa Rica / Richard Condit, Rolando Pérez, Nefertaris Daguerre. p. cm. — (Princeton field guides) Includes bibliographical references and index. ISBN 978-0-691-14707-9 (cloth : alk. paper) — ISBN 978-0-691-14710-9 (pbk. : alk. paper) 1. Trees—Panama—Identification. 2. Trees—Costa Rica—Identification. I. Pérez, Rolando, 1963– II. Daguerre, Nefertaris, 1969–

III. Title.

QK480.P2C66 2011 582.16097287—dc22

2010005197

British Library Cataloging-in-Publication Data is available This book has been composed in ITC Cheltenham with Gill Sans Family Display Printed on acid-free paper. ∞ Printed in the United States of America 10 9

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Contents For ew ord

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Par t I. In t r oduc t Io n c hapter 1. Forests of Panama and c osta r ica c hapter 2. t ree Identification 17 c hapter 3. Species and a reas c overed 23 Pa r t I I. SPec IeS t r eat men t S by Fa mIl y a canthaceae 26 a chariaceae 28 a doxaceae 30 a nacardiaceae 30 a nnonaceae 38 a pocynaceae 62 a raliaceae 76 a recaceae 78 a steraceae 86 betulaceae 86 bignoniaceae 88 bixaceae 94 boraginaceae 96 burseraceae 106 c annabaceae 112 c apparaceae 114 c aricaceae 118 c elastraceae 120 c hloranthaceae 120 c hrysobalanaceae 124 c lethraceae 132 c lusiaceae 132 c ochlospermaceae 144 c ombretaceae 144 c ornaceae 150 c yrillaceae 152 d illeniaceae 152 ebenaceae 154 elaeocarpaceae 154 erythropalaceae 156 erythroxylaceae 158 euphorbiaceae 160 Fabaceae 176 Fabaceae—c aesalpinioideae 176 Fabaceae—mimosoideae 186 Fabaceae—Papilionoideae 214

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Fagaceae 238 Hernandiaceae 238 Humiriaceae 240 Hypericaceae 242 Icacinaceae 246 l acistemataceae 246 l amiaceae 248 l auraceae 252 l ecythidaceae 262 l epidobotryaceae 266 l ythraceae 268 magnoliaceae 268 malpighiaceae 270 malvaceae 272 malvaceae—bombacoideae 274 malvaceae—byttnerioideae 282 malvaceae—Grewioideae 286 malvaceae—malvoideae 292 malvaceae—Sterculioideae 294 malvaceae—t ilioideae 296 melastomataceae 296 meliaceae 310 moraceae 322 muntingiaceae 334 myricaceae 336 myristicaceae 336 myrsinaceae 344 myrtaceae 350 n yctaginaceae 358 o chnaceae 360 Papaveraceae 362 Pellicieraceae 362 Phyllanthaceae 362 Piperaceae 366 Podocarpaceae 368 Polygonaceae 368

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Contents Proteaceae 372 Putranjivaceae 374 r hamnaceae 374 r hizophoraceae 378 r ubiaceae 380 r utaceae 410 s alicaceae 414 s apindaceae 426 s apotaceae 434 scrophulariaceae 446

PArt iii . s u PPo rt in G MAte r iAl Appendix 1. t erminology 467 Appendix 2. Major l eaf t raits of t ree Families Known in Panama and c osta r ica 469 Appendix 3. Families n ot included 475 l it er At u r e c it ed in de x 481

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simaroubaceae 448 siparunaceae 450 staphyleaceae 450 t ernstroemiaceae 452 t hymelaeaceae 452 u lmaceae 454 u rticaceae 454 Verbenaceae 460 Violaceae 460 Vochysiaceae 462

Foreword Forests of the tropics are famous for high species diversity. In Panama and Costa Rica, 200 or more species of trees can be found on a walk of a few hundred meters. This shocks visitors from Europe or North America, where a similar walk might uncover at most 10 species of trees. But those same visitors often study the ground layer of plants in a northern forest, and they are very likely to have learned more than 200 species of smaller plants. Many learned these plants by studying field guides such as the classic by Roger Tory Peterson and Margaret McKenney (1968). Peterson and McKenney’s guide covers far more than 200 species, and thus it provides access for casual botanists to a wide diversity of plants. For me (RC) growing up in North America, these field guides, for both plants and birds (Elias 1980; Petrides 1972; Ridgely 1978), were the basis of my development as an ecologist. It seems very likely to me I would not have ended up a professional in natural history, botany, and ecology if I had not had these sorts of guides to study. When I first took ornithology in Illinois, A Field Guide to the Birds (Peterson 1934) was in constant use on my tables and desks. Later, the Field Guide to Pacific States Wildflowers (Niehaus and Ripper 1976) was also just as important when I first studied botany in California. In contrast, for two of us growing up in Panama (RP, ND), such field guides were scarce, and there were no plant guides. Our botany teachers at the University of Panama captured our attention, but there were no color plates to study over the dinner table, only technical floras (Croat 1978). Many potential botanists of Central America are very likely lost at this career stage because of this gap in their botanical studies. We are trying to remedy the gap by offering an attractive book of photos and range maps for a large number of tree species. Our intent is that budding botanists and ecologists will often have it handy while they become familiar with the plants they encounter near their homes or on walks in the woods. If our impact can be anywhere near the impact of

Peterson’s classics, maybe there will be many more botanists and more such books in coming decades.

Acknowledgments We foremost would like to thank Robin Foster, our mentor in tropical botany and tropical ecology. He first showed us how to study details of leaves to identify trees even when there were no flowers present, traditionally not the way botanists learn. Another prominent contributor to our work has been Suzanne Lao, who keeps track of complex databases and thus helps us organize complicated and oftenchanging scientific names. We also thank the Smithsonian Tropical Research Institute, with its excellent modern facilities in Panama City, for all manner of logistical support for our forays into Central American forests. Both the Smithsonian and the Center for Tropical Forest Science provided us financial support for the work leading to this book. Moreover, the Smithsonian and the University of Panama, especially Professor Mireya Correa, let us consult specimens in their herbarium collections. We would like to thank several scientists for providing access to online databases on Neotropical plants. In particular, the following groups helped us assemble large numbers of tree records: Peter Jørgensen, Trisha Distler, and Chris Freeland (Missouri Botanical Garden); Nick King and Andrea Hahn (Global Biodiversity Information Facility); Barbara Thiers and Anthony Kirchgessner (New York Botanical Garden); and Brian Enquist, Robert Peet, Steven Dolins, Brad Boyle, Peter Jørgensen, Barbara Thiers, Mark Schildauer, Naia Holmes, and Brian McGill (the BIEN Working Group). We also thank Brad Boyle for many comments on early drafts and his plant family descriptions from Costa Rica, and Charlotte Taylor and Susanne Renner for comments on sections covering their specialties. Finally, we are grateful to Salomón Aguilar, Javier Ballesteros, Oldemar Valdés, Victor Galindo, Rolando Pérez, Jr., Nicole Pérez, and Bianeth Pérez for assistance collecting plants and taking photos,

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Foreword

to Barbara Clauson for her careful editing, including Latin names, and to Julia Gonzalez for preparing the index. We dedicate the book to Tobias Schultz, Nat Condit-Schultz, Luc Condit-Schultz, Rolando Pérez, Jr., Nicole Pérez, Bianeth Pérez, Juan Daguerre, and Dionisia Núñez. Richard Condit Rolando Pérez Nefertaris Daguerrre Panama City, Panama 19 September 2009

Part I

Introduction

Chapter 1

Forests of Panama and Costa Rica Tree Diversity in the Tropics of Central America Southern Central America excels in tree species richness. On a world map, Costa Rica and Panama, along with western Colombia, Ecuador, and Peru, have the highest plant diversity (Barthlott et al. 1996). Understanding why begins with understanding the topographic and climatic diversity of the region. The Andes in South America and the central cordillera of Central America are a long, near-continuous spine of mountains, and the variation in elevation from lowland to mountain peaks creates variation in habitat. From the warm lowlands to the cool and windswept mountain tops, communities differ, whether of trees, birds, or butterflies. Particularly important in Central America is the contrast between the Pacific and Caribbean flanks of the mountains, which we will return to shortly. But there is more to high diversity in Central America than a mountain range. Colorado and Switzerland have mountains, but nowhere near the number of tree species. Indeed, all of North America has about 1000 tree species, compared to 2300 in much smaller Panama. We don’t know the total number of tree species in all of South America, but it is somewhere around 20,000. Scientists have debated this contrast in species richness since von Humboldt traveled the Amazon 200 years ago, and there are nearly as many theories as there are theoreticians. One obvious possibility is glaciation: during the past 200,000 years, while North America and Europe were under ice sheets, Central America maintained a moist, warm climate and dense vegetation. There is plenty of evidence that northern regions suffered extinction during the glacial epoch of the past few million years. High species richness is one reason botanists find it difficult to learn, describe, and document plant richness of Latin America. There

are far fewer botanists, more species, and less money available for environmental concerns than there are in North America. Indeed, there remain immense gaps in our knowledge of trees in Central America. We have carried out inventories for 20 years throughout Panama, yet we are still unable to identify every tree we see. In fact, in more remote areas that are difficult to visit, it is typical for tropical botanists to leave as unidentified 25% of the species encountered. There is nowhere in North America where an experienced botanist would have anywhere near this difficulty. Those North American botanists have excellent field guides (e.g., Elias 1980; Petrides 1972). Comparable guides for trees anywhere in the tropics are scarce, though the number has been greatly expanding in the past decade (e.g., Zamora et al. 2000, 2004). Our hope here is to follow this trend and help other biologists expand their knowledge about the tropics. This guide covers 493 tree species, including 438 with color photos, and we offer descriptions of key features for identification and show where they are known in Panama and Costa Rica. We also offer brief notes about the tree species we do not cover, where they are known, and which of the illustrated species they resemble.

Geography, Climate, and Forest Panama and Costa Rica are well inside the tropics, at 8–10°N latitude, where an atmospheric conveyor belt delivers storm systems out of the east—off the Caribbean—during the northern summer. During the winter, from December through April, this belt shifts southward and the region has its annual dry season. The mountain axis of Central America is high enough to intercept storms off the Caribbean, and thus the wettest areas of Panama and Costa Rica are on the Caribbean flanks of the mountains. There, annual precipitation is greater than 3000 mm, with many sites

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getting 4000 or 5000 mm. The Pacific slope of the isthmus is generally drier, getting less than 3000 mm, and as low as 1500 or even 1000 mm at the driest locations. Though generally drier than the Caribbean slope, the Pacific slope is variable and has both wet and dry areas (Figs. 1.1 and 1.2). The central axis of mountains between the Caribbean and Pacific is nearly everywhere wet, often with clouds and fog. To the tree species of Central America, the variation in rainfall is crucial. On the wettest Caribbean slopes, there is enough moisture even during the December–April dry season to keep the soil damp, and forests are wet and green all year. In contrast, much of the Pacific slope has hard, dry soil by April, and many species are deciduous then. By sitting on the cusp of this gradient from dry forest to wet forest, Panama and Costa Rica capture a great diversity of tree species, because the ability to tolerate a dry season is a crucial adaptation. In the 1950s and 1960s, a pioneering ecologist named Leslie Holdridge worked for years in Costa Rica and Panama (Holdridge 1967). Holdridge noticed how the variable climate of the region has a conspicuous impact on the forests and developed a system for classifying vegetation to describe it. The Holdridge system has been widely used by tropical ecologists since, especially in Central America. There are a total of 17 categories in the Holdridge system, but a good basic understanding of forest communities in Central America can be gained by abstracting five broad units: dry, moist, wet, lower montane, and upper montane. (1) Dry forest. In Panama and Costa Rica, dry forest is found in two regions where rainfall is less than 1500 mm annually, one in central Panama west of Panama City, the other in northwestern Costa Rica (Fig. 1.1). True dry forest in the tropics is short, seldom more than 20 m tall, and many trees lose their leaves during the dry season, quite different from the classic image of tropical rainforest. Fine-leaved trees of the legume family are common. In the driest areas of Central America, there are even columnar cacti and the mesquite tree (Prosopis jubata), both common sights in North American deserts. But it is not a desert in Panama and Costa Rica: trees would occupy the entire region, given the chance. We will return to human impacts later.

(2) Wet forest. The lowland forests of the Caribbean slope are wet forests of the Holdridge system. Here, the classic image of rainforest holds: trees are tall, the canopy is dense and continuous except for small gaps created by fallen trees, and epiphytes are conspicuous. Trees do not shed their leaves during the weak dry season. Much less conspicuous is what you can decipher only after learning tree species: wet forests have many more species than do dry forests. There are substantial blocks of wet forest throughout Central America, including extensive and unbroken stretches in San Blas, much of western Panama, and the national parks of Costa Rica’s Caribbean slope as well as the Osa Peninsula (Fig. 1.1). (3) Moist forest. For the forests between wet and dry, we use Holdridge’s phrase “moist forest.” This grows where more than 1500 but less than 3000 mm of rain falls, but where the dry season is pronounced and lasts 3–4 months. Trees are tall; indeed, the biggest trees in the area are characteristic of moist forest (e.g., the kapok [Ceiba pentandra] and the espavé [Anacardium excelsum]). On the other hand, there are deciduous species during the dry season. All the easy-to-see forest near the Panama Canal is moist forest (Fig. 1.2), as is a large block in Darien National Park in eastern Panama and much of the Pacific slope of Costa Rica (Fig. 1.1). The remaining climatic zones in our abbreviated version of Holdridge are montane: forests above 800 m elevation. There is an important division at about 1500 m, so we consider two categories. (4) Lower montane forest. From about 800 to 1500 m elevation, forests resemble lowland wet forests, with tall trees and dense canopy, and wet and lower montane zones share many species. Tropical cloud forests fall in this range, and are typical on the tops of lower mountains, where clouds often sit. There are high densities of epiphytes—orchids, bromeliads, ferns, and mosses—in cloud forests. (5) Upper montane forest. Above 1500 and especially 2000 m elevation, forests are quite different, and are sometimes classified as temperate, like forests of North America. Indeed, oak (genus Quercus) and alder (Alnus) are conspicuous above 2000 m, along with many other temperate groups familiar in North America

Forests of Panama and Costa Rica

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Santa Rosa NP

Caribbean

Braulio Carrillo NP Guanacaste NP San Jose

La Amistad Intl Park San Blas Nusagandi

Bocas del Toro El Cope Cerro Boquete Punta

Corcovado NP Osa Peninsula

Chiriqui

Santa Fe

Darien

Sarigua NP Herrera Los Santos Azuero Peninsula

Pacific Ocean

Figure 1.1. Place names and forest zones in Panama and Costa Rica. The dry zone is indicated by yellow, the moist zone by light green, and the wet zone by dark green. A light green topographic line demarcates the lower montane zone, at 800 m elevation, and a black line shows the upper montane zone, at 1500 m.

Caribbean Fort Sherman

Santa Rita Ridge

Upper Chagres River

Colon

Fort San Lorenzo Chagres R.

Soberania NP Pipeline Rd Barro Colorado I.

Cerro Jefe

Madden Dam Gamboa Sendero El Charco Panama Canal

Cerro Azul

Summit Garden

Parque Metropolitano

Panama City

El Valle Punta Chame

Altos de Campana NP

Pacific Ocean Figure 1.2. Place names and forest zones in the Panama Canal Area. The dry zone is indicated by yellow, the moist zone by light green, and the wet zone by dark green. The only topographic line is at 800 m elevation and demarcates the lower montane zone (there is no elevation above 1500 m in the canal area).

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and Europe. Among these temperate trees, though, is a high diversity of tropical taxa. Upper montane forests are not giant forests. Trees are generally less than 20 m tall, and on windswept mountain peaks, trees become sparse or even absent. The boundaries between these zones are blurred, and the use of 5 (or even 17) categories is arbitrary. The transition between all types is gradual. There is no place one can stand and point to one forest type on the left and another on the right. Three of the five zones we describe are especially blurred: moist, wet, and lower montane. But categorization is how humans learn to understand variation, and this is why the Holdridge system became popular. To aid in understanding the climate zones, Figure 1.3 shows three distribution maps illustrating typical tree species ranges that are linked to forest zones. Where forest remains, all five forest zones are rich in tree species by the standards of North America or Europe, though wet forest is richer than dry forest. Crucial to Central American diversity is another kind of diversity—the change in species between climatic divisions. The wet Caribbean slope is very different in tree species from the moist and dry forests of the Pacific half of the isthmus, and montane forests differ from either, though they are much closer to wet forests. Near the Canal in Panama, the wet forests of Santa Rita and the moist forests near Panama City (Fig. 1.2) are only 50 km apart but share almost no tree species. Much of our scientific research has been about this sharp change in tree species composition in central Panama and the rainfall patterns that cause it (Condit et al. 2001, 2002, 2004, 2005; Engelbrecht et al. 2007). In Costa Rica, very different species can be seen within 150 km in Santa Rosa National Park in the northwest to Braulio Carrillo in the northeast, crossing the mountains in between (Fig. 1.2).

Human Impact As we all understand too well, forests are altered and often simply removed by humans, and the categories of Holdridge do not include human impacts. In fact, deforestation is closely linked to climatic zones (Condit et al. 2001). Humans prefer the tropical dry climate to the

tropical wet climate, and tropical dry forests have largely been removed in Central America and indeed much of the world. In Panama and Costa Rica, the drier Pacific slope is mostly pasture interspersed with cities and towns, and we don’t really know what mature dry forest would look like, because there is none. In contrast, the mountains and the Caribbean lowlands of Central America still retain much of their forest and have far fewer humans. Tellingly, there is substantial Pacific forest in wet areas, such as the west side of the Azuero Peninsula in Panama and the Osa Peninsula in Costa Rica (Fig. 1.2), as well as on mountains near the Pacific, where the climate is wet due to high elevation.

Visiting Forests in Panama and Costa Rica Both countries have lovely and extensive forests readily accessible via good roads. In Panama, the easiest forests to reach—and the most familiar to us—are moist lowland forests near the Canal, from Panama City to Gamboa and Pipeline Road (Fig. 1.2). Parque Metropolitano near the city has hiking trails, and though the forest has been heavily disturbed, it is rich in tree species. Taller and more attractive forest can be found at Sendero El Charco, just past Summit Gardens, on the main highway toward Gamboa, and there is a short hiking trail there. Both Metropolitano and El Charco are in moist, secondary forest, with many species typical of the Pacific half of the isthmus. The paved road ends at Gamboa, which is surrounded by moist secondary forest. Just beyond starts Pipeline Road, a long dirt road through unbroken forest in Soberania National Park. The first half of the road has a recent disturbance history, but beyond the Rio Limbo, the forests are mature and have been little impacted for at least a century. From a tree perspective, Pipeline Road provides a fascinating switch from the Pacific to the Caribbean: the first two kilometers have a Pacific coast element, with Bursera simaruba and Castilla elastica, but the stretch of road up the steep hill after the Rio Limbo has species of Caribbean coast wet forest (e.g., Welfia regia is easy to see). Permission and a fourwheel-drive vehicle are needed to reach the

Lozania pittieri

Forests of Panama and Costa Rica

15

Lozania pittieri

Caesalpinia coriaria

Caesalpinia coriaria

Alnus acuminata

Alnus acuminata

Figure 1.3. Sample species distribution maps illustrating major forest types. Lozania pittieri (Lacistemataceae) has a typical wet-forest distribution, mostly on the Caribbean half of the isthmus, but conspicuous at the Osa Peninsula and other Pacific sites where rainfall is higher. It also occurs at some lower montane sites. Caesalpinia coriaria (Fabaceae—Caesalpinioideae) is only found in the two dry zones of central Panama and northwestern Costa Rica. Alnus acuminata (Betulaceae) is confined to the highest mountains in far western Panama through Costa Rica. Red dots are locations from herbarium records or our own inventories, where precise coordinates were noted. All provinces where a species is known are also stippled; because there are records where the province was noted without coordinates, there may be stippling in areas with no dots. Moreover, provinces are political boundaries and not climatic zones, and stippling in a province does not necessarily mean the species occurs throughout the province. Notice that the map of A. acuminata shows stippling along the Caribbean coast in Costa Rica, even though the tree only occurs in the central mountains. This is because boundaries of several provinces straddle the mountains and reach the coast.

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Chapter 1

wet-forest area, but it is an excellent area to visit for both flora and fauna. The best example of undisturbed tropical moist forest in Panama is on Barro Colorado Island. It is intermediate between the coasts geographically and also floristically, and species of the Pacific slope (such as the cuipo [Cavanillesia platanifolia]) along with hints of wetter forests (Virola surinamensis as a large example) are common there. Barro Colorado Island also has the biggest trees in Panama, all either kapok (Ceiba pentandra), espavé (Anacardium excelsum), or the sandbox tree (Hura crepitans). The island can be visited with permission from the Smithsonian Tropical Research Institute. Excellent Caribbean wet forest also can be visited in a 1-day trip from Panama City, though not as easily as Pipeline Road or forests near the city. A road runs up the Santa Rita Ridge from the Transisthmian Highway, and not far from the highway, near the radio towers, there are small stands of true Caribbean wet forest. Further up the road there are more extensive stands of forest, but it is difficult driving, requiring four-wheel drive and even then not recommendable after wet weather. Similar wet forest can be visited at Nusagandi, about 4 hours east of Panama City, where there are rustic accommodations for tourists. Both Santa Rita and Nusagandi are highly diverse and claim many wet-forest species impossible to see at Barro Colorado Island or Pipeline Road. Another site near the Caribbean that can be reached from Panama City in a few hours is San Lorenzo National Park, at the mouth of the Chagres River. There are accommodations at the former US military base called Fort Sherman. Montane forest at 800–1000 m in elevation is also easy to visit near Panama City, at Cerro Campana about 1 hour west of Panama City, and Cerro Azul and Cerro Jefe, which are just north of the Tocumen (Panama City) airport. Cerro Campana has good trails through cloud forest, and there are various trails and cabins around Cerro Jefe. Both sites are much wetter than the nearby lowlands and have some cloud-forest specialists as well as a broad contingent of wet-forest species that otherwise occur only near the Caribbean coast. There are several other areas to visit mon-

tane forest in Panama: El Valle not far from Cerro Campana, El Cope, and several sites in Chiriqui, including La Amistad International Park. There is a road from the Pan-American Highway over the mountains to Bocas del Toro that crosses through cloud forest near the Fortuna Dam. Further west, the popular sites for visitors are around the towns of Boquete and Cerro Punta, and this is the only area in Panama where forests above 2000 m in elevation can easily be visited. These harbor a temperate-like flora, with oaks, alder, dogwood, and elm, but also many tropical taxa. Unfortunately, nearly all true dry forest in Panama has been cut down. Forested sites near Panama City are drier than the Caribbean coast but are really moist forest. Savanna-like remnants of dry forest can be seen along the Pan-American Highway west of Cerro Campana and at Sarigua National Park near the town of Chitre, but you need to visit Costa Rica to see extensive dry forest. Costa Rica has excellent parks spanning the range of climates, indeed, Panama’s parks aspire to match Costa Rica’s system. Near San Jose, the capital city, Santa Rosa, Guanacaste, and Braulio Carrillo national parks offer easy access and excellent trails. Santa Rosa, in the far northwest, has been allowed to regenerate for 30 years, and is the only spot in either Panama or Costa Rica where true dry forest can be seen. From Santa Rosa eastward over the mountains and down through Braulio Carrillo offers a marvelous transect through all the climate zones. In the far south, the Osa Peninsula is peculiar for having a wet-forest climate near the Pacific coast. The extensive forest there is protected in Corcovado National Park, and there are lodges around the park and remote camps and cabins within. Indeed, thanks at least partly to the popularity of ecotourism, visitors can easily reach all manner of forests in Costa Rica and Panama without much difficulty, from remote and undisturbed wildernesses to city parks. Whether by bus, rented automobile, or with tour groups, all forest zones of the two countries can be visited within a day of the major cities. We hope this book will help nature tourists learn something about the trees as excellent guides (Ridgely 1978) do for the birds.

Chapter 2

Tree Identification Learning Plant Families We emphasize the importance of learning the traits of plant families as a step in identifying species. Families are groups of related species that usually share conspicuous and easy-tolearn characteristics. In animals, dogs (wolves, foxes, etc.) are a family and cats (leopards, bobcats, etc.) are another. In both examples, few people have any difficulty recognizing the similarities that link groups of species within the families or any trouble separating dogs from cats. Plant families are likewise readily distinguished, though sometimes not as obviously as dogs and cats. Once you know families well, the list of potential matches for the specimen in your hand is greatly narrowed. A second reason to learn plant families is that the number of entities is manageable. We cover 83 plant families in this book, a reasonable number to keep track of. Since learning half the families would make you an expert on Central American flora, you have much to gain from knowing 40-odd groups of plants. Consider it otherwise: there are 2300 tree species in Panama, and to be an expert you should probably know 600 to 800. That is too many entities for most of us to memorize. Learning families is emphasized in Gentry’s (1996) A Field Guide to the Families and Genera of Woody Plants of Northwest South America. We used this book extensively in writing our family descriptions and compiling tables of traits, and anyone set on learning tropical trees of the Americas needs to study Gentry’s book. In our family descriptions, we also relied heavily on Flowering Plants of the Neotropics (Smith et al. 2004). As a warning to botanists from North America or Europe, though, the families of trees that dominate Costa Rica and Panama are nearly all unfamiliar in northern forests. Apart from palms, a visiting botanist without tropical experience will find nothing recognizable. Maple,

beech, hickory, pine, fir, and redwood—very common trees of forests across the United States and Canada—are not found in the tropics at all. Instead, the major tree groups include the mahogany family (Meliaceae), the coffee family (Rubiaceae), the legumes (Fabaceae), and many more that are either completely unknown or very rare in North America and Europe. To help learn the families and their traits, Appendix 1 is a brief glossary of terms used here and elsewhere in the book, and Table 2.1 and Appendix 2 offer a tabular summary of families and their characters. We present drawings in the next section, and other plant guides (Johnson and More 2004; Petrides 1972) have excellent drawings as well. Even better, Harris and Harris (1994) have written a great book on plant vocabulary with drawings, and Wikipedia has a thorough Glossary of Botanical Terms.1

The Major Leaf Characters The starting point in learning plant families and thus identifying tropical tree species is to know the six basic leaf characteristics. In most cases, these characteristics are easy to see. Much as a good bird-watcher first checks for the bill size of an unidentified bird, these are the traits you first check when examining an unknown tree: (1) opposite versus alternate leaves; (2) simple versus compound leaves; (3) toothed or lobed leaf margins; (4) latex; (5) clustered versus regular spacing of leaves; (6) stipules 1 http://en.wikipedia.org/wiki/glossary_of_botanical _terms.

18

Chapter 2

Table 2.1

Leaf Characters of 25 Common Tropical Tree Families of Central America

Leaf

Family

Alternatesimple

Anacardiaceae* Annonaceae Apocynaceae* Boraginaceae Chrysobalanaceae Euphorbiaceae Lauraceae Lecythidaceae Malvaceae* Moraceae Myristicaceae Rutaceae* Salicaceae Sapotaceae Urticaceae Violaceae* Apocynaceae* Clusiaceae Melastomataceae Myrtaceae Rubiaceae Violaceae* Anacardiaceae* Burseraceae Fabaceae Malvaceae* Meliaceae Rutaceae* Sapindaceae Bignoniaceae

Oppositesimple

Alternatecompound

Oppositecompound

Latex

Stipule

Toothed

Lobed

Two-ranked

Compound form

yes drips drips

yes yes

some many

drips yes

yes yes

many some yes

yes

most

yes yes

many

yes

drips most yes

drips yes some yes yes

many some

yes yes yes some

odd odd odd, even palmate odd, even odd odd palmate, bipinnate

Families are divided into four groups based on the two fundamental characters (alternate versus opposite leaves, and simple versus compound leaves). Families with an asterisk appear twice. “Some,” “most,” and “many” refer to the proportion of species in the family; “yes” indicates all species (or nearly all). Under latex, “yes” means latex appears but does not drip. “Odd” and “even” refer to odd- and even-pinnate, respectively.

The first two leaf traits are the most important and have to be learned. These are always the first two characters to check. (1) Opposite versus alternate leaves. Opposite means that two leaves arise from the same position on the branch, so a pair of leaves are precisely opposite one another (Fig. 2.1, bottom). Conversely, alternate means that each leaf arises by itself (Fig. 2.1, top). Note that the terminology might lead to confusion. In both diagrams of Figure 2.1, leaves are on opposite sides of the branch. The determining feature is whether two leaves arise from one spot on the branch (opposite) versus singly (alternate). In most species, whether leaves are opposite or alternate is unambiguous, but there are questionable cases. One confusing arrangement is called subopposite, where two leaves

are nearly but not exactly opposite. Another ambiguity is where leaves are so closely crowded that it is difficult to see whether they are opposite or alternate. Also, there is a third though rare possibility, with more than two leaves arising from a single spot. These are whorled leaves. (2) Simple versus compound leaves. Compound means that individual leaves are divided into separate sections called leaflets. Simple leaves are not thus divided. In the example in Figure 2.2, one leaf has five leaflets. To many who have never studied botany, each leaflet would just be called a leaf, so the structure in Figure 2.2 would be a branch with five leaves. With just a little experience, it is nearly always easy to tell simple from compound.

Tree Identification

19

leaflet

rachis

alternate leaves

petiole

Figure 2.1. Alternate versus opposite leaves.

stipule

pulvinus opposite leaves

There are two tricks. First, the compound leaves on one tree have approximately the same number of leaflets. This means that the units of leaf tissue tend to come in groups of about the same size. But the “approximate” qualifier is important: a tree might have most leaves divided into five leaflets, but some will have three or seven. Count how many leaflike units there are in several different groups: if the leaves are not compound, then you are counting the number of leaves per branch, and this number may vary a great deal. But if the leaves are compound, you are counting leaflets per leaf, and the number will be consistent, usually varying by less than four. The next trick for establishing whether leaves are compound is to check the spot where the petiole, which is the stalk carrying the leaf, emerges from the branch (Fig. 2.2). In compound leaves, this is nearly always swollen relative to the rest of the petiole and is called a pulvinus. If you are looking at a branch with five leaves, there will not be a swelling. Once you have determined that leaves are compound, there are a wealth of obvious features that help identify the tree. The first and most obvious is the number of leaflets, which, as we just noted, is fairly consistent within species. Leaves with just two (bifoliate) or three (trifoliate) leaflets are very consistent (Fig. 2.3, bottom), hence the special names for these cases. With more leaflets, the consistency is looser, so that trees with 21 leaflets are likely

Figure 2.2. A compound leaf with five leaflets.

to also have leaves with as few as 17 up to 25 leaflets, even on the same branch. A second obvious trait of compound leaves is leaflet arrangement. Most compound leaves are pinnate, meaning the leaflets are arranged in two rows on opposite sides of the rachis (Fig. 2.2). Pinnate leaves can have either an odd number of leaflets, or an even number (Fig. 2.3), and this is very consistent within species: in any one tree, you will see odd-pinnate leaves or even-pinnate leaves, but not both.2 Generally, odd-pinnate leaves have a series of leaflet pairs plus one terminal leaflet, whereas even-pinnate leaves lack the unpaired terminal leaflet (Fig. 2.3). Given that leaves are pinnate, another simple character to check for is whether leaflets are opposite or alternate along the rachis.The definition is the same as for alternate and opposite leaves, but applied now to leaflets: two leaflets emerging opposite one another, from the same position on the rachis, are called opposite leaflets, but if leaflets arise singly they are alternate along the rachis. In either situation, the number of leaflets may be even or odd. As a caution, note that the 2 There are botanical terms, imparipinnate (odd) and paripinnate (even), but these are good examples of unnecessary technical jargon, because odd- and even- are far easier to remember (see Appendix 1).

20

Chapter 2 leaflet

pinnule

leaflet

leaflet

odd-pinnate

even-pinnate

bipinnate

bifoliate

trifoliate

palmate

Figure 2.3. Various types of compound leaves. In each, the entire structure is a single leaf, that is, there are six leaves shown in this figure. The most common type is on the far left, where odd refers to the number of leaflets.

presence of a terminal leaflet is not always obvious when leaflets are alternate (Fig. 2.4). Other arrangements are less common. When leaflets radiate from a single point, like spokes of a wheel (Fig. 2.3), the leaves are called palmate because the arrangement resembles a hand.3 Leaves can also be bipinnate, which means that each leaf is divided once into leaflets, and each leaflet divided again into “subleaflets.” The precise, botanical term for “subleaflet” is pinnule (Fig. 2.3). Determining whether a leaf is bipinnate requires applying the rules described above—counting leaflets and pinnules and checking for a swollen petiole base—twice. (You may be wondering whether further divisions are possible, and indeed, one of the 493 species we describe is tripinnate.) Having determined whether the leaves are simple or compound, you may need to reconsider whether they are opposite or alternate. Study Figure 2.4. Whether leaves are alternate or opposite hinges on the arrangement of the entire leaf: in (a) and (b) in the figure, the leaves are alternate, whereas in (c) and (d), they are opposite. It is thus possible (and not unusual) for leaves to be alternate while leaflets are opposite, and Figure 2.4 shows other possibilities. Throughout the species descriptions we will repeatedly apply the alternate– opposite dichotomy to leaves and separately to leaflets—check the pictures carefully. (3) Toothed or lobed leaf margins. This is 3

The term digitate also is used (see Appendix 1).

easy to see and may be noticed before the first two traits listed above. Oaks are famous for lobed leaves, and every Canadian is familiar with the maple leaf’s three main lobes. Teeth are finer, like those on elm leaves. Lobes are always conspicuous, but teeth can be sparse or inconspicuous, and sometimes a wavy margin can be mistaken for teeth. Leaf teeth are also less consistent within families than leaf arrangement (alternate–opposite) or leaf division (simple–compound), that is, quite a few families have some species with teeth and some without. This is why this character comes third and not first. (4) Latex. Break off a fresh leaf and check the broken end. Most plants lack latex, and no liquid will appear. But in some species, liquid will drip out rapidly, and in others, droplets will appear at the wound but will not drip. In most cases, latex is conspicuous, but there are species that produce small amounts of barely visible (sometimes clear) latex. Generally, all parts of the tree, not just the leaf, exude latex, so if you break a branch or cut deeply into the bark, you will see some. This is why many trees in well-utilized tropical forests have machete slashes—someone was checking for latex to confirm an identification. Another important detail is that latex dries up. If you pluck a leaf in the field but carry it home to study, the latex may no longer be there. It is a character you should note when the leaf is fresh. (5) Clustered versus regular spacing of leaves. Clustering may seem a vague feature,

Tree Identification

a

c

21

b

d

Figure 2.4. Leaf and leaflet arrangements in pinnate leaves. a) Odd-pinnate leaves with nine leaflets. Leaves are alternate, leaflets are opposite. b) Even-pinnate leaves with six leaflets. Leaves are alternate, leaflets are alternate. c) Odd-pinnate leaves with nine leaflets. Leaves are opposite, leaflets are opposite. d) Even-pinnate leaves with six leaflets. Leaves are opposite, leaflets are alternate.

but it is surprisingly consistent within plant families and thus very useful. At one extreme, leaves are spaced very regularly along the branch, and at the other, leaves are clustered toward the branch tip. In the former case, all the leaves typically lie in a flat plane; if you remove such a branch, it will rest easily on flat ground. These are called distichous or tworanked leaves. In contrast, clustered leaves are usually oriented at many angles, and a branch broken free will not rest flat on the ground. Leaf arrangement can be questionable, for instance if leaves are only slightly bunched, and of course the arrangement is obscured when leaves fall off. But leaf arrangement is always a character worth examining. (6) Stipules. These are the subtlest of the

six main traits: you know you are a botanist when you study stipules. They are tiny, weak appendages where the petiole meets the branch (Figs. 2.2 and 2.5), and they frequently fall off leaving only an inconspicuous scar. Many families lack stipules altogether, and though tiny, stipules come in a variety of forms that create family-identifying features. Indeed, stipules are valuable traits in some of the most difficult groups to learn. A problem you will have, though, is knowing when the absence of stipules means they fell off or were never there. Always look for stipules first at the outermost ends of branches, where they are most likely to be retained, then check for thin scars at the base of older leaves, where stipules would have once been.

22

Chapter 2

stipule

If you spend any time at all trying to identify tropical trees, you will get to know these six major traits and their terminology well, and you will be noting each in every species you see. Soon, you will be counting leaflets and breaking off leaves to look for latex in ornamental plants at shopping malls. There are, of course, many other traits that help identify tropical trees, including trunk traits (overall size, buttresses, bark, and branch arrangement), leaf traits (venation, color, hairs, size, odor, and petiole length), plus flowers and fruits. You should be wondering why we omitted these other features from our master list, and there are reasons. The six main traits are (usually) strong dichotomies, easy to observe, and they tend to distinguish families well. Most other traits fail in one respect or another: they represent subtle gradations (leaf color or leaf size), or they do not distinguish families (bark or hairs). Many of these other traits will come up in our descriptions of individual species, but they are less useful at the family level. What about flowers and fruits? In botanical keys, they usually come first, and indeed, they are usually more characteristic of families and species than the leaves. The unfortunate truth is, however, that during any visit to a forest anywhere in the world, you will find that very few individuals are reproductive. If you rely on flowers or fruits, you will identify almost nothing. We include photos of flowers for most species, but you must focus on leaves, branches, and the trunk if you want to make any progress identifying trees.

Figure 2.5. Stipule and stipule scar. At the lower pair of leaves, the stipule has fallen, leaving a scar behind (a line across the branch). These are opposite leaves, but alternate leaves have stipules and scars as well, as do compound leaves (Fig. 2.2).

Categorizing Families by Major Leaf Traits Table 2.1 summarizes the six major leaf traits for 25 of the most common tropical tree families, not including the palms (Arecaceae), which are easy to recognize without studying traits. The simplest use of the table is to help memorize the six major opposite-leaved families and the seven major compound-leaved families. Species of these families are fairly easy to learn. But note the large number of families with simple, alternate leaves. This is the prototypical tropical tree, and telling those families apart requires delving into more subtle traits, such as leaf clustering and stipules. Even experienced botanists struggle with the large number of species with simple, alternate leaves. Knowing the 25 major families well is a great start, but the diverse tropics have many more families than these 25, and in Appendix 2 we tabulate the important characters of 117 families, which cover just about every family of tree you will ever encounter in Panama or Costa Rica. In this full list, there are 78 families that include species with alternate, simple leaves. Even considering that 16 of those are very seldom seen (marked “scarce” in Appendix 2), and 9 of the rest are strictly montane, there are still 53 families you have to consider if you come across simple, alternate leaves in lowland tropical forest (we cover 46 of those in the main text). This allows us to reiterate the importance of recognizing compound and opposite leaves, because the number of families with either or both those traits is much smaller.

Chapter 3

Species and Areas Covered This is a guide to the trees of Panama and Costa Rica. Both the species and the regions covered, however, require some explanation. First, our own experience has been in Panama, based at the Smithsonian Institution’s tropical research station at Barro Colorado Island, an island in the Panama Canal, and at offices in Panama City. We know Panama much better than Costa Rica. But the two countries share 72% of their tree species, so our photographs and descriptions work well in both regions. We have spent 15 years compiling a tree species checklist for Panama. There are still holes and questions, but at this moment we know 2321 tree species native to Panama. Our main sources have been the flora of Panama by Correa et al. (2004), plus online lists of herbarium specimens at the Missouri Botanical Garden,1 the Global Biodiversity Information Facility (GBIF),2 and the New York Botanical Garden.3 We have not yet compiled a tree species list for Costa Rica, but based on occurrence records in the latter two databases, 1680 of the 2321 are known in Costa Rica. Assuming Costa Rica has the same number of total species as Panama, the two countries combined would have almost 3000 species. We have not tracked down those species in Costa Rica not known in Panama, but some can be found in the excellent yet still incomplete series on Costa Rican trees (Holdridge and Poveda 1975; Zamora et al. 2000, 2004). Based on the Missouri Botanical Garden and GBIF online databases, we created range maps for all 481 native species in this book, covering both Costa Rica and Panama (sample maps are shown in Fig. 1.3). The maps show indiTropicos Web database, http://www.tropicos.org/. Global Biodiversity Information Facility data downloads from Panama and Costa Rica, http://data.gbif.org/ countries/browse/. 3 http://www.nybg.org. 1 2

vidual locations where an herbarium specimen was collected, if the exact coordinates were recorded. Provinces where a species was collected are also stippled, including cases where exact coordinates were not recorded but the province was (which is why you will find provinces shaded even without dots within). The maps are good by tropical standards, but there remain many gaps and we made no attempt to fill them: if a species is known at one end of Costa Rica and the opposite end of Panama, there is a good chance it will eventually be found in between. Perhaps our maps will help spur botanists to fill the gaps. Astute readers will notice a few errors too, most obvious where a tree occurs in the ocean. (The databases of the Missouri Botanical Garden and GBIF have millions of records, and it is currently not plausible to catch all the errors.) An additional difficulty in a guide about tree species is establishing the criteria that define a tree. Our definition of a tree includes any plant with a woody stem that can grow taller than 2 m and stands on its own. Omitted are woody vines, or lianas, which spend most of their lives leaning on other trees. Included are woody plants 2–4 m tall that we refer to as treelets, because many would prefer not to call such small plants trees. Excluded are woody plants always less than 2 m tall. Our biggest trouble assembling the checklist of Panama’s trees and treelets was the minimum size criterion. The tropics are rich in species that grow in the forest understory and look like little trees, but that are usually categorized as shrubs or even subshrubs. Look in particular at our discussions of the genera Ardisia (Myrsinaceae), Miconia (Melastomataceae), and Psychotria (Rubiaceae). All three have many species that grow to a height very near our limit of 2 m. Because few people ever measure the exact height of trees or treelets, we were left making educated guesses about which of those species to include in the tree

24

Chapter 3

checklist. If we were to slightly broaden our definition to include even shorter treelets, we would have to add about 40 Psychotria species; narrow the definition to include only trees greater than 4 m tall, and we would lose 40 Psychotria. We decided to exclude both lianas (woody vines) and epiphytes (trees growing on other trees). This creates additional uncertainty because some species grow as free-standing trees on some occasions and as lianas on others; likewise for epiphytes. Moreover, there are plenty of species we have never seen firsthand, and there are a number of species for which we could not find clear descriptions of the mode of growth. By far the larger problem we had in deciding what to include in the book was the limited printing space available confronting more than 2300 known species. To make a reasonable-sized book with good photos and descriptions meant space for 493 species, with 438 illustrated. We deliberately chose the most numerous and widespread species, based on our own observations. Especially near the Panama Canal, you will find that a high fraction of individuals are covered in this book. There will be additional cases where you should be able to find the genus, but not the species. For all families and genera, we include brief comments on all species known in Panama but not described or illustrated. This is a great help when attempting to identify an unknown specimen that is not an obvious match to photos in the book. In Appendix 3, we list the tree families known in Panama and Costa Rica but not covered in the main text, and Appendix 2 lists the major leaf traits for nearly all the families.

A different sort of difficulty in preparing the book comes from the plant names. We rely largely on Latin names, as most in the tropics do, though we include many local names in Spanish. Those local names, however, are not well known and vary from region to region. You will need to rely on Latin names to be certain about identifications, but Latin names add another sort of difficulty: synonyms, or cases where two different scientific names are used for the same plant. Scientists have precise rules about Latin names, indeed, only one name can be the officially accepted name. However, names change, meaning that the accepted name in a publication from the 1980s may not match the name in current use. Many beginning botanists find this very frustrating. In this book, we always use the most recent accepted name according to Correa et al. (2004) and the Missouri Botanical Garden.4 In the Index, we include all synonyms we consider relevant for every species. The descriptions that follow are organized alphabetically by family, and within family alphabetically by genus then species. As explained at length in Chapter 2, organizing by family is important for learning tree species. Nevertheless, many species can be learned even without knowledge of the families, and you may sometimes leaf through this book without paying attention to families. Indeed, once you are familiar with the flora of a region, you will identify many species at a glance without thinking of the family. We hope our photos and descriptions get you closer to that stage.

4

Tropicos Web database, http://www.tropicos.org.

Part II

Species Treatments by Family

26

Acanthacea

Acanthaceae

(Including Avicenniaceae)

Aphelandra sinclairiana

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A large family found throughout the world, with 3200 species worldwide and 1700 in tropical America. Most are herbs or small shrubs, but a very few are trees. Many species in the group are cultivated as ornamentals for their showy flowers and are found in gardens nearly everywhere. Panama has 12 species we designate as trees, but this depends considerably on how “trees” are defined: the genera Aphelandra and Ruellia have 50 species between them, only a few of which qualify as treelets. We cover three species here, including two mangroves. Acanthaceae are best known for their showy flowers, which are usually long, tubular, and brightly colored. They are shaped like mint flowers, with two lower lips at the opening of the tube. Leaves are nearly always opposite and often clustered at branch ends. Of the three species we cover, however, only the first matches this general family description well;

the two mangroves have to be learned separately. See also the mint family, Lamiaceae, and the Verbenaceae, which are similar and related, indeed, several species have recently been shifted among these families.

Aphelandra sinclairiana. A treelet of forest understory. Leaves are opposite, untoothed, clustered near the top of the stem. Flowers are unmistakable, composed of long pink tubes with extended lower lips, held on stalks that have orange bracts between the flowers; before the flowers open, the stalk of bracts looks like a shrimp’s tail. Distribution: In forests of the Caribbean half of the isthmus throughout Panama and Costa Rica, from lowland to lower montane. Fairly common near Pipeline Rd., where it often grows in groups and is very conspicuous when flowering. Recognition: Mostly known from its flowers and overlooked without them. With leaves alone, Acanthaceae should remind you of Rubiaceae, but lack the stipule scar. There are three more Aphelandra species in Panama that we count as treelets; all are small, have weak stems, and have long, tubular pink or orange flowers. There are quite a few more species in the genus that are too small to count as treelets.

Avicennia bicolor

Avicennia bicolor

Avicennia bicolor (mangle salado, mangle negro,

mangle prieto, black mangrove). A tall tree of salt marshes. See description of the next species, which is very similar, but A. bicolor has gray to white leaf undersides. Distribution: Along the coast in muddy swamps, only known on the Pacific of c. Panama and nw. Costa Rica, where it overlaps with A. germinans.

Acanthacea

Aphelandra sinclairiana

Avicennia bicolor

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28

Achariaceae

Avicennia germinans

Avicennia germinans

Avicennia germinans (mangle salado, mangle

negro, mangle prieto, black mangrove). A tall tree of salt marshes. The trunk is straight and cylindrical, without stilts or buttresses of any sort. The soil surface surrounding the trunk typically has pneumatophores extending above the water, roots that breath while the soil is submerged. Their connection to the main tree is hidden beneath the mud, so the pneumatophores look like strange aquatic fungi. Bark is dark, divided into rectangular plates by vertical and horizontal fissures. Leaves are opposite, shiny green, narrowing at the base, on a short petiole. Fruit is an oval capsule, covered with spongy material that floats, then splits open to release seeds. Distribution: Only grows in seawater swamps, on

Achariaceae Lindackeria laurina

both Pacific and Caribbean coasts, though more widely on the Pacific. It is usually the dominant mangrove further from the sea than Rhizophora, that is, Rhizophora is the one at the margin of the ocean, with Avicennia behind it. Recognition: The mangrove habit is found in unrelated tree species in several families. Besides the closely related A. bicolor, there are four other principal mangroves to compare: Laguncularia and Conocarpus (Combretaceae), Rhizophora (Rhizophoraceae), and Pelliciera (Pellicieraceae). Rhizophora and Pelliciera have very distinct trunks, and Conocarpus has alternate leaves. Laguncularia is most like Avicennia, but its leaves have small glands at the base. There are five more genera of Acanthaceae we count as trees: Barleria, Bravaisia, Pseuderanthemum, Ruellia, and Trichanthera. All have a single species except Ruellia, which has two. Trichanthera is a small tree of forest edge fairly common in the Canal Area, with purple flowers; it is not known in Costa Rica. Both Avicennia and Trichanthera were formerly in the Verbenaceae, and Avicennia is sometimes placed in its own family, Avicenniaceae.

(Formerly Flacourtiaceae)

Lindackeria laurina

family is not easy to recognize from the leaves, but the one common Panama species is easy to learn due to an odd quirk.

Lindackeria laurina (carbonero). A small or me-

A small tropical family of mostly shrubs and trees, with 145 species worldwide and about 40 in the Americas. The members we treat here used to be classified in the now-defunct family Flacourtiaceae. (One group of the old flacourts is now joined with the Achariaceae, the other with the Salicaceae.) Panama has five species of the newly expanded Achariaceae, of which we cover two and illustrate one. Achariaceae leaves are alternate, simple, and lack latex. In many species, the leaves are held perpendicular to the petiole, and are clustered irregularly. In many Achariaceae, leaves are toothed, but not all, and the one species we illustrate lacks teeth. The

dium-sized tree of forest edge in drier areas. When mature, the trunk has many stilts at the base. Leaves are large and wide, rounded at the base, and bunched toward branch ends. They hang downward at a sharp angle to the long upright petioles. Most distinctive is what happens when a leaf is crumpled: white lines immediately appear at all the folds. Flowers are white, in dense groups on branches below the leaves. Fruits are green, spiny capsules. Distribution: A common secondary-forest species near the Canal, principally on the drier Pacific slope, often along roadsides in wooded areas. Scarce in mature forest. Common and easy to see at Pipeline Rd. Otherwise known sparsely throughout Panama and Costa Rica. Recognition: When first learning the species, fold the leaves and you will always know it. The way the big leaves hang downward from long petioles is obvious after some experience.

Achariaceae

Avicennia germinans

Lindackeria laurina

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Adoxaceae

Mayna odorata

Mayna odorata

Mayna odorata. Not illustrated. A small wet-forest tree with narrow, slightly toothed leaves held at an angle to the petiole; the petiole is swollen at both ends. Flowers are large, white, in tight groups on branches just below leaves. Fruits are spiny green

Adoxaceae

Viburnum costaricanum

balls just like those of Lindackeria. Distribution: Widely collected in lower montane and Caribbean coast sites in e. Panama, including Santa Rita and Cerro Campana. Also known on the Osa Peninsula in Costa Rica. Recognition: The toothed, bunched leaves and petioles are fairly distinctive. Another Mayna species is known from the Caribbean slope but it is not known in Costa Rica. Another genus of Achariaceae, Carpotroche, has two species, both similar in leaf form to M. odorata, but Carpotroche has a larger fruit growing off the branches and lacking spines. One of the two is widespread on the Caribbean slope of Costa Rica and Panama, the other sparsely known and only in c. Panama.

(formerly Caprifoliaceae)

Viburnum costaricanum

A family of 450 small trees and vines, almost all in the temperate Northern Hemisphere. This is the family of elderberry, honeysuckle, and Viburnum, all of which have many well-known and often cultivated northern species. There are several tropical Adoxaceae whose ranges extend south through the mountains of Central and South America. The four species in Panama are in high-elevation forests; we cover one here. Species of the family in Central America all have opposite leaves, some with teeth, and can reliably be distinguished by a tiny detail: small, yellow glands on the leaf margin, visible only with magnification.

Viburnum costaricanum. A medium-sized tree of high mountains. Trunk is straight and cylindrical.

Leaves are small, opposite, untoothed, and have very few secondary veins. There are clusters of small hairs on the leaf underside where the veins meet (you will need a hand lens to see). Flowers are white, in flattopped clusters, present much of the year. Fruits are small berries. Distribution: Only above 2000 m elevation in w. Panama and Costa Rica, where it grows in open areas. Recognition: The genus Viburnum is common in North American forests, and is often planted in gardens and yards, and the flat-topped flower clusters of tropical species are typical of the genus. But most northern species have toothed leaves, so the toothless Central American species may be difficult to recognize. Cornus disciflora (Cornaceae) is similar, but Viburnum has fewer secondary veins that do not arch forward as in Cornus. The two genera have similar flowers that can easily be confused. Viburnum has two more species of high mountains in Costa Rica and Panama, both similar but less common. The other genus of Adoxaceae is Sambucus, the elderberry, which has opposite, compound, toothed leaves and is also known only at high elevation.

Anacardiaceae Anacardium excelsum

Anacardium excelsum

A moderate-sized family, with 700 species worldwide and 170 in tropical America. Most are trees or shrubs,

but there are a few lianas; most are tropical, but there are temperate groups, including the North American sumacs (Rhus) and poison ivy (Toxicodendron). In the tropics, the family is best known for the mango and the cashew, both described below. Anacardiaceae are found in most tropical forests, and usually have one or more fairly common species. In Panama, there are 11 species native, and we illustrate 7 and describe 2 more, including the nonnative mango. Anacardiaceae have alternate leaves, and most species worldwide have compound leaves, but some are simple, and the common Panamanian species

Anacardiaceae

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Viburnum costaricanum

Anacardium excelsum

fall in both categories. When compound, Anacardiaceae leaves are odd-pinnate, with a single terminal leaflet and the rest in opposite pairs. The base of the petiole, where it meets the branch, is swollen, brown, and flattened or grooved (not cylindrical). Several families share this collection of traits, most importantly the Burseraceae. Experienced botanists distinguish the Anacardiaceae by a faint resinous

smell from broken leaves, light and pleasant, but it can be very faint and takes practice to be useful. Burseraceae have a similar odor, but usually more pronounced. Once learned, the odor is very useful, because both simple- and compound-leaved anacards have it. Regardless, the common species of Anacardiaceae in Panama are not difficult to learn, because all have fairly distinctive leaves or bark.

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Anacardiaceae

Anacardium excelsum (espavé, wild cashew). A huge and conspicuous tree throughout Panama. There are no buttresses, but the base of the trunk is usually swollen and irregular. Leaves are simple, alternate, long and narrow, widest near the end, densely bunched at the end of branches; older leaves have yellow veins toward the tip. Leaves on juvenile plants are much longer than those on adults. On the ground beneath big trees in the forest, there are usually great piles of blackish leaves. Flowers are small, greenish white, in loose terminal clusters. The fruit is shaped like a kidney, resembling the cashew. Distribution: One of the most abundant trees in the Canal Area and much of the Pacific slope of both Panama and Costa Rica. Not seen on the Caribbean slope except near the Canal. Often abundant along streams in dry areas and in open areas and secondary forests; around Panama City it forms nearly pure stands. Less common in mature forest, where giant trees occur sparseAnacardium occidentale

Anacardium occidentale

Anacardium occidentale (cashew, marañón). An abundant small and crooked tree of farmland. Usually branches close to the ground. Foliage is dense throughout the crown, reaching almost to the ground. Leaves are short and oval, and main veins are yellow. Most trees have a few red leaves. Flowers Astronium graveolens

Astronium graveolens

Astronium graveolens (zorro, ron-ron, tigrillo,

tolerante, cucaracho). A tall canopy tree with a straight cylindrical trunk, moderately buttressed when large. The bark is dark gray, but peels off in pieces to reveal large, light-colored circles of inner bark. Leaves are alternate, odd-compound, with 11–15 leaflets that are pointed at the tip, somewhat toothed (conspicuous on juveniles), slightly asymmetric at the base; leaflet pairs are opposite one another. The secondary veins branch near the leaf border, forming a clear Y. Some individuals drop all their leaves toward the end of the dry season, and most drop some

ly but widely. It does not germinate in the shade, so there are seldom juveniles in mature forest. Recognition: No other huge tree is as abundant, so it is easy to learn its largish leaves, large trunk, dark bark, lack of buttresses, and (in the forest) piles of black leaves under the crown. In urban areas and farmland, it can be common as a juvenile and confused with a few other species. See Anacardium occidentale and two species in the Apocynaceae: the ornamental frangipani (Plumeria) and especially its wild relative Laxoplumeria, both with very similar leaves, but dripping white latex when broken. In mature forest, juvenile Gustavia superba (Lecythidaceae) and Alseis blackiana (Rubiaceae) can be confused with espavé, because they also have long, bunched leaves; espavé leaves can usually be distinguished by the yellowish upper veins, and remember also the faint turpentine odor. Campnosperma panamensis looks like espavé, but is only known on the Caribbean coast.

are tiny, in loose clusters above the leaves, with small, pink to greenish petals. The fruit is green, shaped like a kidney, with the edible nut attached to a larger fleshy portion. Distribution: Only in farmland and around human habitation, often planted, but also grows on its own; never seen in forest. Native to the Neotropics, but its natural origin is obscured by centuries of human association. Recognition: Compare to Anacardium excelsum, which has similar though longer and narrower leaves. The most important difference between the two is trunk form—the espavé grows tall and straight, whereas the cashew is short and leaning. In grasslands of dry regions, look for Curatella americana (Dilleniaceae), quite similar to A. occidentale but with sandpapery leaves that do not turn red while on the tree.

leaves. Prior to falling, leaves are bright scarlet, and even in the middle of the wet season, there are typically a few red leaves in the crown. Flowers are small, green to yellow. As fruits develop, five brown sepals remain below in a star form. Distribution: A tree of the drier parts of Panama and Costa Rica; common in mature and secondary forest on the Pacific slope, and fairly common in farmland. Also in mature forest, though not common, plus lower montane sites near the Pacific. Not known on the Caribbean slope. Recognition: Mosquitoxylum, Spondias, and Tapirira (all anacards), plus Bursera (Burseraceae), some Trichilia (Meliaceae), and Zanthoxylum (Rutaceae) have similar odd-compounds leaves with many leaflets. Compare especially Spondias and Astronium, which are very similar. The Y-form of the secondary veins and asymmetric leaflet bases are subtle but solid traits for Astronium, and Spondias has different fruits. In adults, the splotchy bark of Astronium is distinctive; see Beilschmiedia in the Lauraceae, with reddish splotches but much different leaves. Remember also to look for scarlet leaves in big Astronium.

Anacardiaceae

Anacardium occidentale

Astronium graveolens

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Anacardiaceae

Campnosperma panamense

Campnosperma panamense

Campnosperma panamense (orey). Not illustrated. A tall tree of Caribbean coastal swamps. Leaves are very similar to those of the cashew, but the trunk is straight and tall. Distribution: Only occurs in swamps near the ocean in Bocas del Toro and nearby Costa Rica, but is abundant there and forms nearly pure stands. The leaves resemble those of the cashew, but it is a tall, straight-trunked, and

fairly large tree. Recognition: Distinguished by its habitat.

Mangifera indica (mango). Not illustrated. Native to India. A small or medium-sized tree with a short trunk that often branches near the ground. Leaves are long and narrow, dark green, alternate and simple, clustered at the end of branches, and very dense over the entire crown. Flowers are small, in clusters above the leaves. Fruits are unmistakable. Distribution: Farmland and cities everywhere in the world’s tropics. Occasionally occurs in forest in Panama, but only at abandoned farm sites or where humans have brought it. Recognition: The crowns densely packed with leaves are easy to pick out in farmland even when not in fruit. On those rare occasions when it is found in the forest, the mango is not so easy to identify and might pass for Lauraceae or Sapotaceae.

Mosquitoxylum jamaicense

Mosquitoxylum jamaicense

Mosquitoxylum jamaicense. A rare wet-forest

tree. Leaves are alternate, odd-compound, with many

leaflets. The stalk of the terminal leaflet has a slight wing on either side. Juveniles’ leaflets are pubescent. Flowers are small, white, in dense terminal clusters. Distribution: In lower montane sites and wet forest of the Caribbean slope throughout Panama and Costa Rica, but sparse. Some saplings can be seen along the outer section of Pipeline Rd., and it is fairly common between Cerro Azul and Cerro Jefe. Recognition: See Spondias, which is very similar. As juveniles, fuzzy and more numerous leaflets help distinguish Mosquitoxylum, and the slight wings on the terminal leaflet stalk clinch it.

Spondias mombin

Spondias mombin

Spondias mombin (jobo). One of the most wide-

spread and common trees of the region. In farmland, it is a medium-sized and often leaning tree, but in the forest it can be tall and straight. The leaves are alternate, but bunched toward the end of the branches, emanating like spokes of a wheel. They are long and odd-compound, with opposite leaflets. Crushed leaves smell faintly turpentine-like. The

bark is gray, and sometimes (not always) has blunt, wartlike spines; bigger trees lack warts. The small, white flowers appear in dense, pyramidal bunches just after new leaves grow out (late dry season, early wet season). The fruits are yellow, sweet, and edible. Distribution: Abundant in farmlands and towns throughout Panama and Costa Rica, and often used in living fences. It is also common in secondary forests, where juveniles occur only in natural clearings; giant trees occur sporadically in old growth. Recognition: Bursera (Burseraceae) is also common in farmland and has similar leaves but nothing like the warty, gray bark of Spondias. In mature forest where the jobo is not common, it can be difficult to recognize. See first the very similar Spondias radlkoferi and Astronium graveolens, then check Mosquitoxylum jamaicense, Trichilia (Meliaceae), and Zanthoxylum (Rutaceae) for similar, odd-compound leaves.

Anacardiaceae

Mosquitoxylum jamaicense

Spondias mombin

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36

Anacardiaceae Spondias radlkoferi

Spondias radlkoferi

Spondias radlkoferi. A Spondias of the forest. Similar in all respects to S. mombin, but has smooth, gray bark with vertical fissures, lacking warts. Distribution: Widespread but much less common than S. mombin; on both slopes near the Canal, but only on the Pacific slope in Costa Rica. Not usually encoun-

tered outside the forest. It is most common in secondary forest and juveniles only appear in forest clearings. Recognition: These two Spondias species are best distinguished by bark when trees are larger; the smooth, gray bark of S. radlkoferi is quite unlike the rough, warty bark of S. mombin. Fruits differ slightly: those of S. radlkoferi are 3–4 cm in diameter, those of S. mombin are 2–3 cm, and those of S. radlkoferi are green or black at maturity, not yellow. The leaves of the two species are very similar and probably not safely separable. See S. mombin for other similar species. Two other Spondias species are known in Panama. One of them, S. dulcis, was recently discovered by our surveys near the Canal. The other is cultivated and seen widely but is probably not native.

Tapirira guianensis

Tapirira guianensis

Tapirira guianensis (caobilla). A tall forest tree with a straight trunk and medium-sized buttresses. Leaves are alternate, odd-compound, with fewer and somewhat larger leaflets than the other anacards. The terminal leaflet is often angled upward relative to the rest of the leaf. Broken leaves have a strong resinous smell. In dark, wet forests, seedlings of Tapirira can be abundant on the forest floor beneath big trees. Flowers are tiny, yellowish green. Distribution: A major canopy element in wetter forests of the Caribbean slope near the Canal; very common in mature forest in Soberania. Known on the Caribbean slope and the Osa Peninsula in Costa Rica. Recognition: Fewer and larger leaflets with a clear odor make this species resemble the Burseraceae

more than the other Anacardiaceae, and it is perhaps more likely to be confused with Protium costaricense or maybe Tetragastris (both Burseraceae) than with Spondias, though it is similar to the latter. The upturned terminal leaflet is a good character in most trees. Except for Mosquitoxylum, which has more leaflets, all the anacards and Tetragastris are in dry and moist forest, whereas Tapirira is a wet-forest tree; P. costaricense is also known in wet forest, though. A second Tapirira species, T. mexicana, is restricted to mountains of w. Panama and Costa Rica. It has similar compound leaves. T. guianensis has been collected in the mountains, so the two ranges may overlap. We have not mentioned two genera in the Anacardiaceae, Mauria and Toxicodendron, both with a single species and both restricted to mountains of w. Panama and Costa Rica. Mauria has compound leaves with few, large leaflets. Toxicodendron is a small tree with toxic resins just like poison ivy and poison oak of North America, but its long compound leaves are more like those of Spondias than the trifoliate North American species.

Anacardiaceae

Spondias radlkoferi

Tapirira guianensis

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Annonaceae

Annonaceae Anaxagorea panamensis

Anaxagorea panamensis

One of the major plant families in lowland forests throughout the tropics, with 2500 species worldwide and 900 in tropical America. Nearly all are trees, and several are widely planted as fruit crops. Panama has 76 species in its checklist, and we illustrate 26 and describe 2 more. Annonaceae leaves are always simple and alternate, never toothed. They are spaced regularly along branches in a flat plane; leaves of this sort are called two-ranked or distichous. In most Annonaceae, the branches zigzag from leaf to leaf, but sometimes this is not especially evident. Branches are usually squarely perpendicular to the trunk and spaced regularly. In all these traits, Annonaceae are similar to the related Myristicaceae. The two families also share a faint, slightly spicy odor, as do Lauraceae, but this is often too faint to be helpful; it takes a lot of experience to rely on it. Another trait specialists use in Annonaceae is bark toughness: when breaking a twig, bark peels off in long, fibrous strips, and twigs can be difficult to break. This separates the Myristicaceae, which have easily broken twigs and bark. Finally, flowers and fruits of Annonaceae are distinctive. Flowers are on branches, between leaves, and usually have just three thickened petals that are nearly always white or greenish. Fruits are often large, and many have spines or warts. Unfortunately, the consistency of these traits defining Annonaceae work against separating spe-

cies within the family. Indeed, the main genera have similar leaves, and learning them requires knowing each species’ individual traits. Fruits identify genera more consistently, but are seldom present. Without reproductive parts, any tree with two-ranked leaves and neither teeth nor latex has a good chance of being Annonaceae, and zigzag branches strengthen the possibility. But be aware that there are many genera in other families with two-ranked leaves, and some have zigzag branches.

Anaxagorea panamensis. A mature forest tree-

let that grows in dense aggregations. The smallest Annonaceae around, usually with a leaning stem and low branches. Leaves are narrow and pointed as Annonaceae go, shiny and smooth, clearly two-ranked, and zigzagging. Distribution: The most remarkable geographic range of any tree in Panama. We know it from two locations, one on Barro Colorado I. and one near Pipeline Rd. In both spots, there are several hundred small trees in such a dense patch that they are difficult to walk through, and easily visible as a dark dome of treelets from 50 m away. Outside these patches, there are none at all—we have never seen an isolated individual. It grows in similar patches in Costa Rica. Overall, known from only a few sites on the Caribbean slope of both countries. Recognition: The two-ranked leaves match the Annonaceae pattern; as such, the small size of the tree and narrow and pointed leaves should suggest this genus. The habit of growing in discrete and very dense clumps is a good identifier. Three more species of Anaxagorea are found in Panama. All are treelets of forest understory. A. allenii is known in lowland wet forests east of the Canal, much more widespread than A. panamensis, but not known in Costa Rica. The other two species are rarely seen in Panama, but are widespread in wet forests of Costa Rica.

Annona glabra

Annona glabra

Annona glabra (anon de agua, anon de puerco,

anon de pantano, guanábana de pantano, pond ap-

ple). A small tree of ponds and marshes. Leaves are simple, alternate, but not obviously two-ranked as in most Annonaceae. Secondary veins are inconspicuous. Flowers are cream-colored outside, pink inside. Fruits are large, smooth, like an apple (hence the English name). Distribution: Near the Caribbean coast at river mouths and swampy areas and around Gatun Lake in Panama. In Costa Rica, seen along both coasts and near several lakes. Its roots are often in the water. Recognition: Best known by its location at the water’s edge and the big fruits, which are often present.

Annonaceae

Anaxagorea panamensis

Annona glabra

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40

Annonaceae

Annona hayesii

Annona hayesii

Annona hayesii (canelo). A medium-sized forest

the two-ranked pattern of the family. Veins are deeply impressed and secondary veins conspicuous. Distribution: Known only near the Canal to e. Panama. Widespread in forests on the Pacific side of the Canal Area, and especially abundant near Panama City. Recognition: The deeply impressed veins make this quite easy to recognize, given the annonaceous leaf arrangement. The nonforest A. purpurea has the same style venation, though, but notice that the fruits are completely different.

tree. Leaves are fairly large and broad, and fit closely Annona pittieri

Annona pittieri

Annona pittieri (anon de montaña). A small tree of montane forests. The leaves are shiny and strongly aromatic. Flowers are produced out of the main trunk or the largest branches. Distribution: Lower

montane and lowland wet forests throughout Panama and Costa Rica, but sparse. However, fairly common at Cerro Campana. Recognition: A. montana (not pictured) is very similar; but is largely a Caribbean wet-forest species, whereas A. pittieri is also found in the mountains (there may be places where they co-occur). Both those species, plus the cultivated A. squamosa (not pictured) have big, warty fruits, but A. squamosa does not occur in the forest, and fruits of A. montana have spines. Several other Annonaceae have shiny leaves: Oxandra with smaller leaves; Xylopia with inconspicuous venation; Duguetia confusa with a yellowish tint on the leaf underside.

Annonaceae

Annona hayesii

Annona pittieri

41

42

Annonaceae

Annona purpurea

Annona purpurea

Annona purpurea (toreta). A small or medium-

sized tree of dry, open areas, sometimes cultivated.

Leaves are big, with veins impressed above, and have a modestly strong aroma when broken. Flowers are brownish on the outside, but reddish purple within. Fruits are large and warty. Distribution: Mostly on the Pacific slope of c. and w. Panama and Costa Rica, in dry areas. Common in secondary forests, including Parque Metropolitano in Panama City. Recognition: The leaves are like those of A. hayesii, with prominently impressed veins. A. spraguei also has prominent veins, but the leaves are bluish underneath; Guatteria amplifolia is similar, but with less-impressed veins and much different fruits.

Annona spraguei

Annona spraguei

Annona spraguei (chirimoya, negrito). A medi-

um-sized tree of roadsides and edge. Leaves are long, narrow for their length compared to other Annona, two-ranked, with branches zigzagging between leaves. Veins are conspicuous above; secondary veins are straight and parallel, fairly densely spaced. The leaf underside has a tint of light bluegreen due to a dense coat of fine hairs. Flowers are solitary, greenish yellow outside and purple inside. The fruits have long, thin, wormlike green spines forming a dense covering. Distribution: Only c. to e. Panama. One of the most common roadside trees around Panama City. Inside the forest, occurs only in

natural clearings. Recognition: The modestly large leaves with closely spaced, parallel secondary veins and a bluish cast on the underside are characteristic. Rollinia pittieri also has such a bluish cast, but secondary veins are much different, and it occurs only in wet forests. In the roadside range of A. spraguei, Castilla elastica (Moraceae) has leaves with similar size and shape but lacking the closely spaced secondary veins and the bluish color; Castilla exudes white latex from a broken leaf. See A. hayesii and A. purpurea, whose leaves are somewhat like those of A. spraguei, but lack the bluish undersides. Virola species (Myristicaceae) have bluish leaf undersides, and V. surinamensis could be confused with A. spraguei; however, it is found exclusively in mature, wet forests. One other Annona species, A. acuminata, is native, and it is common at Barro Colorado I. and nearby. It is a treelet with small, narrow leaves. There are six more Annona species, mostly growing in rural areas, and their natural distributions are obscured by frequent cultivation. They have large, warty fruits.

Annonaceae

Annona purpurea

Annona spraguei

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44

Annonaceae

Cymbopetalum lanugipetalum

Cymbopetalum lanugipetalum

Cymbopetalum lanugipetalum (yayito). A

small tree of wet forests. The trunk is small and often branched not far above the ground. Leaves are simple, alternate, two-ranked, pointed at the tip, fairly narrow, and have a corrugated look due to im-

pressed secondary and tertiary veins. Distribution: Wet and lower montane forests of the Caribbean slope of c. and e. Panama, plus wet forests of sw. Costa Rica. Common in the Fort Sherman area at the Caribbean mouth of the Canal. Recognition: The corrugated leaves are easy to spot and distinctive; however, Unonopsis sp. nov. of Caribbean wet forests with similar leaves also is very common around Fort Sherman. Three other Cymbopetalum species are known in Panama. One is restricted to the Darien, and the other two are rare in Panama but fairly widely known in wet forests of Costa Rica. These others lack the well-impressed veins of C. lanugipetalum.

Desmopsis panamensis

Desmopsis panamensis

Desmopsis panamensis (yayito). An understory

treelet of mature forest. Trunk is straight and unbranched. Leaves are alternate and two-ranked, shiny and smooth above, veins inconspicuous. A small, brown, fuzzy appendage, about 3 mm long, is found at the base of each leaf and lays pressed against the upper leaf surface. Flowers are yellowish, on long stalks. Fruits are small berries in clusters. Distribution: Known at only a few locations from

Costa Rica to e. Panama, and mostly rare, but with a major concentration at Barro Colorado I. and Pipeline Rd. in c. Panama, where it is one of the most abundant small trees. It only occurs inside mature forest, never on forest edge or open areas. Recognition: The leaves are arranged as in other Annonaceae, but the smooth surface with inconspicuous venation is not typical for the family. The little brown, pointed flap lying on the leaf base is distinctive and always present. Five other Desmopsis species are known in Panama. Four are montane and found from c. Panama through Costa Rica; one of those is also found throughout the Caribbean lowlands of Costa Rica. The fifth species is very poorly known. Desmopsis usually has an odd floral trait: the long stalk has a small leafy appendage near the base. Without flowers, leaves are not easy to characterize.

Annonaceae

Cymbopetalum lanugipetalum

Desmopsis panamensis

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Annonaceae Duguetia confusa

Duguetia confusa

Duguetia confusa. A medium-sized tree of wet

forests. The trunk usually has a low fork, and the bark is black. Leaves are simple, alternate, regularly spaced, two-ranked, with a light, slightly yellowish

underside due to fine yellow scales; the scales also impart a rough feel to the leaf. Flowers are creamcolored, and the fruit is baseball-sized and warty. Distribution: Wet and lower montane sites throughout. Recognition: The leaves are very typical Annonaceae in size and shape, but the yellow-green underside is not typical for the family; see also Guatteria aberrans. Four other Duguetia species are known in Panama, but three are known only from far e. Panama. The fourth, D. panamensis, is known widely but sparsely on the Caribbean slope of Panama and Costa Rica; it also has yellowish scales on the leaf underside.

Guatteria aberrans

Guatteria aberrans

held on a long stalk. In general, Guatteria is a difficult genus if there are no fruits or flowers. Panama has 18 species; we treat 5.

Guatteria aberrans (sigua negro). A large, wet-

Genus Guatteria. The biggest South American

genus of Annonaceae and by far the dominant one in mature forest. Indeed, in the understory of wet forests in the American tropics, Guatteria is a good guess for any unknown Annonaceae. They have the typical annonaceous pattern of regularly spaced leaves held in a flat plane. Most have moderately impressed veins on the upper surface, not as deeply as Annona hayesii, though. Leaves of Eschweilera (Lecythidaceae) are similar to those of Guatteria. Flowers are on short stalks from branches, but have various colors and shapes. Fruits are much different from those of Annona, consisting of a cluster of small berries on short (sometimes very short) stalks, resembling Desmopsis, but in the latter, the cluster is

forest tree. The trunk is straight, and large trees have small buttresses. Bark is black. Leaves are in flat plane, alternate, regularly spaced along zigzag twigs. The veins above are not impressed as in other members of the genus. Leaf underside has short pubescence and is much lighter in color than the upper side. Secondary veins are parallel, closely spaced, and conspicuous. Distribution: Wet forests in c. Panama only; fairly common at Santa Rita, also on mountains around Cerro Jefe. Recognition: The leaf arrangement follows the family-wide pattern well, but the lack of impressed veins detracts from its resemblance to other Guatteria. The closely spaced secondary veins are reminiscent of Annona spraguei, which also has a light-colored leaf underside. The black trunk of G. aberrans resembles that of G. dumetorum. In addition, G. recurvisepala and Duguetia confusa are similar, and Virola (Myristicaceae) might also be confused.

Annonaceae

Duguetia confusa

Guatteria aberrans

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Annonaceae

Guatteria amplifolia

Guatteria amplifolia

Guatteria amplifolia (yayito). A small tree of mature forest. Leaves are alternate, regularly

spaced, in a flat plane. Veins are deeply impressed, and the leaves are large, with a very short petiole. Distribution: On the Caribbean slope throughout, plus the Osa Peninsula and a few lower montane sites. Common in the Canal Area, from Barro Colorado I. to the Caribbean and nearby mountains. Recognition: The large size of conspicuously bullate leaves makes for fairly easy recognition, given the annonaceous pattern. Two Annona with deep leaf creases, A. purpurea and A. hayesii, are fairly similar, but have smaller leaves.

Guatteria dumetorum

Guatteria dumetorum

Guatteria dumetorum (sigua negro). A large forest tree. The trunk is cylindrical or slightly fluted; big trees broaden at the base into small buttresses. The

bark is dark or even black. Leaves are alternate and arranged regularly, in a flat plane, and veins are modestly impressed. Distribution: Wet areas from c. Panama to w. Costa Rica. Very common and widespread in mature forests near the Canal, and one of the major canopy species in the old forest at Barro Colorado I. Juveniles are common and easy to find inside the forest. Not seen in open areas. Recognition: Where big trees are common, learning the black and slightly buttressed trunks is not difficult. The leaves are very typical of the family and the genus. Beware the similar though unrelated Eschweilera pittieri (Lecythidaceae).

Annonaceae

Guatteria amplifolia

Guatteria dumetorum

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50

Annonaceae

Guatteria jefensis

Guatteria jefensis

Guatteria jefensis. Not illustrated. A small tree of the Cerro Jefe area. Leaves are dark above, light yellow below, regularly spaced along zigzag branches, but rotated from the branch so not in a flat plane. Flowers are solitary, on long stalks off branches below leaves, and have rounded, green petals. Fruits are like those of other Guatteria. Distribution: A true narrow endemic, known only from Cerro Jefe. Recognition: Branches look very typical of Annonaceae, but the rotated leaves are unusual for the family. Range overlaps that of G. sessilicarpa, but that species has stiff leaves with revolute edges.

Guatteria recurvisepala

Guatteria recurvisepala

Guatteria recurvisepala (yayito). A mediumsized wet-forest tree. Leaves are alternate, spaced

regularly and in a flat plane, pointed at the tip; branches zigzag from leaf to leaf. Veins are impressed, but not greatly so. The leaves have reddish hairs on the veins, especially conspicuous on the underside, and the tip of the twig and terminal bud are also pubescent; all are rough to the touch. Distribution: Mostly grows at lower montane sites, but also in lowland wet forests, though sporadic. Recognition: The reddish hairs are a good character, because few Annonaceae are so conspicuously covered (but G. aeruginosa, not illustrated and common in Costa Rica, has similar hairs).

Guatteria sessilicarpa

Guatteria sessilicarpa

Guatteria sessilicarpa. A medium-sized wet-for-

est tree. Leaves are alternate, spaced regularly and in a flat plane; branches zigzag from leaf to leaf. Compared to other Guatteria, leaves are smooth and shiny above, with veins only slightly impressed. Leaves are fairly narrow and pointed at both ends, and the edg-

es curve downward slightly (revolute). Distribution: Another narrow endemic, this species was first described from the Santa Rita ridge east of Colon, and is only known from there, Cerro Jefe, and Nusagandi in San Blas. It can be found fairly easily around Cerro Jefe. Recognition: The zigzag branches and flat leaf plane indicate the family, but otherwise, the smooth, revolute leaves do not immediately bring Guatteria to mind. The 13 Guatteria species not described are all from wet or lower montane forest, very sparsely known in Panama, though G. aeruginosa, G. diospyroides, and G. oliviformis are more common in Costa Rica. It will take careful study to separate them.

Annonaceae

Guatteria recurvisepala

Guatteria sessilicarpa

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Annonaceae

Mosannona garwoodii

Mosannona garwoodii

Mosannona garwoodii. A small, understory tree restricted to the Canal Area. Leaves are shiny above, narrow and pointed, widely and regularly spaced along zigzag branches. The leaf underside is a remarkable sky blue color, covered in a soft pubes-

cence (indeed, it is the hairs that give the color). Distribution: Discovered on Barro Colorado I. and only named recently (for botanist Nancy Garwood). We have seen it elsewhere on the Caribbean side of the Canal Area and east of Madden Lake. It is strictly in shady forest interior. Recognition: No other species around has truly sky blue leaf undersides; Xylopia and Virola (Myristicaceae) are bluish, but not so deeply so. Leaf shape also resembles that of Xylopia, but veins in all Xylopia are inconspicuous to the point of being barely visible; in Mosannona, veins are clearly visible both above and below. Two more Mosannona species are restricted to far e. Panama and are seldom seen.

Oxandra longipetala

Oxandra longipetala

Oxandra longipetala. An understory treelet. The small trunk forks near the ground and sometimes has small stilts at the base. Leaves are simple,

alternate, regularly spaced and in a flat plane, shiny and lustrous. The petiole is very short, and the leaf itself short, almost square in shape. Flowers are white. Distribution: Abundant in the understory of mature forest at Pipeline Rd.—by far the dominant species there. Also common around Fort Sherman near the Caribbean Canal entrance. Seldom seen elsewhere. Recognition: Leaves have the same shape and size as those of Licania hypoleuca (Chrysobalanaceae), but leaves of Licania are white below. Oxandra is very easy to learn around the Limbo Hunt Club at Pipeline Rd., which is the only place you are likely to see it.

Annonaceae

Mosannona garwoodii

Oxandra longipetala

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Annonaceae

Oxandra panamensis

Oxandra panamensis

Oxandra panamensis (malagueto de montaña).

A small tree of forest understory. The trunk is short and branches near the ground. The bark often peels off. Leaves are shiny above, narrow, short, and pointed, regularly and tightly spaced along zigzag branch-

es. The leaf underside is clear green, only a bit lighter than above; veins are very inconspicuous above and below. Distribution: In the Canal Area, seen nearly exclusively on limestone soils near Madden Dam and Fort Sherman; seldom seen elsewhere. Where it occurs, it can be very common. Recognition: Narrow, pointed, shiny leaves are much like those of Xylopia, but darker and more lustrous above and lacking the blue-green underside of Xylopia. See also Anaxagorea and Mosannona. One more Oxandra species occurs in the area, but it is very rare and only found on the Pacific coast of w. Panama and s. Costa Rica.

Rollinia mucosa

Rollinia mucosa

Rollinia mucosa (annonillo, chirimoya). A medi-

um-sized tree of wet and cloud forests. Leaves are fairly large, oblong, arranged in a flat plane and regularly spaced along the branches. Veins are widely

spaced and conspicuous, but not impressed. Tips of the twigs are slightly hairy. Flowers have three fleshy petals, and fruits are large, round, and covered with warty spines. Distribution: Mainly a tree of lower montane and cloud forests throughout, but also occurs in some low-elevation wet forests. There are a few along Pipeline Rd. Common at Cerro Campana. Recognition: The genus Rollinia is related to Annona, and has similar large fruits. This species and the next are much like A. spraguei, A. hayesii, and A. purpurea. Without fruits, Rollinia is somewhat like Guatteria aberrans and could easily be confused with other Annonaceae.

Annonaceae

Oxandra panamensis

Rollinia mucosa

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Annonaceae Rollinia pittieri

Rollinia pittieri

Rollinia pittieri (annonillo). A medium-sized tree

of wet forests. Quite like the previous species in most respects, but tips of twigs lack hairs, leaves are lighter colored below, flower petals are not twisted, and fruits are smaller and on long stalks. Distribution: Widespread in Caribbean lowland forests and lower mountains. Recognition: See the previous species and several similar Annona. The two Rollinia have different ranges, with R. pittieri common at lower elevations, but they may overlap.

Unonopsis panamensis

Unonopsis panamensis

Unonopsis panamensis. An understory tree of wet forests. Leaves are two-ranked and regularly spaced, very long and narrow, veins not much impressed. Flowers are tiny and grow off the trunk. Distribution: Known very sparsely in wet lowlands of c. Panama to Costa Rica. Recognition: Perhaps you are finding the Annonaceae difficult to tell apart; rest assured, professionals do too. A variety of herbarium specimens take this name, almost certainly more than one species.

Annonaceae

Rollinia pittieri

Unonopsis panamensis

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58

Annonaceae

Unonopsis pittieri

Unonopsis pittieri

Unonopsis pittieri (yaya). A medium-sized forest tree. Leaves are typical Annonaceae, tworanked and regularly spaced, with veins modestly impressed, fairly narrow and pointed but less so than in other Unonopsis. Flowers are very small. Fruits a small cluster of berries, first green but turning red, on branches below leaves. Distribution: In wet to lower montane forest from c. Panama through Costa Rica. Reaching moist forest near the Canal, where it is fairly common in Barro Colorado I. old growth and at Pipeline Rd., inside the forest. Recognition: Leaves are not especially distinct, re-

sembling those of Desmopsis rather than other Unonopsis.

Unonopsis sp. nov. An understory wet-forest treelet that we believe is an undescribed species. Leaves are two-ranked and regularly-spaced, very long and narrow, veins deeply impressed. Distribution: Fairly common near Caribbean in the Canal Area. Recognition: Easy to recognize due to the very long, corrugated leaves. It has been misidentified as U. panamensis, which also has long leaves but lacks the impressed veins. Naucleopsis naga (Moraceae) can have long, narrow leaves, but is fig-like and should not be confused. There is a rare genus in the Ulmaceae, Ampelocera (not pictured), having one species with long narrow leaves, but it does not have impressed veins. Eleven more Unonopsis names are used in Panama. Generally, they have rather nondescript Annonaceae leaves and fruits like Guatteria; flowers are typically very small.

Annonaceae

Unonopsis pittieri

Unonopsis sp. nov.

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Annonaceae

Xylopia aromatica

Xylopia aromatica

Xylopia aromatica (malagueto hembra). A common and easy-to-recognize roadside tree. Branches are long and hang downward, giving the tree a conical form, like a Christmas tree. Leaves are narrow, pointed, shiny above and bluish below, secondary

veins so faint they are almost invisible. As Annonaceae go, leaves are crowded along the branches, but regularly spaced and in a flat plane, as typical for the family. Flowers are white, with long, narrow petals, produced singly on branches among the leaves. Distribution: A very common roadside and farmland tree, in disturbed areas and secondary forest, throughout the Pacific slope. Seldom inside the forest; only in natural clearings. Recognition: This species should never be confused because of its distinctive branching pattern and small, narrow leaves. X. frutescens also is a conical, roadside tree, but with smaller leaves. The remaining Xylopia grow in the forest.

Xylopia bocatorena

Xylopia bocatorena

Xylopia bocatorena. Not illustrated. A mediumsized tree of wet forests near the Caribbean. Leaves

and branches are very similar to those of X. macrantha. But X. bocatorena often has newly grown reddish leaves dangling downward from branches. Flowers have short, wide petals, unlike X. aromatica and X. frutescens, which have narrow petals. Distribution: Abundant in deep forest shade of lowland wet forests, more widespread and common than X. macrantha. Does not occur in moist forest of the center of the isthmus (absent from Barro Colorado I. and Soberania). Recognition: Often has some red leaves, providing a useful character; X. macrantha does not have red leaves.

Xylopia frutescens

Xylopia frutescens

Xylopia frutescens (malagueto macho). A com-

mon and easy-to-recognize roadside tree. Branches ascend from the trunk, then angle downward, im-

parting a Christmas-tree–like appearance. Leaves are narrow, pointed, very small, and from a distance might resemble needles of a conifer (leaves of Podocarpus [Podocarpaceae] are similar). Distribution: A very common roadside and farmland tree, in disturbed areas and secondary forest, throughout the Pacific slope of Panama and Costa Rica, but also in wet and even lower montane forests. Usually not inside the forest. Recognition: Much like X. aromatica in form and roadside habitat, but with much smaller leaves. The leaves resemble those of X. sericea, but the latter is entirely a wet-forest species.

Annonaceae

Xylopia aromatica

Xylopia frutescens

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Apocynaceae

Xylopia macrantha

Xylopia macrantha

Xylopia macrantha (malagueto de montaña). A medium-sized tree of forest interior. The dark trunk

has white lenticels. Branches are horizontal. Leaves are just like those of X. aromatica. Flowers have thick, cream-colored petals. Distribution: This is mainly a species of moist to wet forests of the Caribbean slope. Common in the old forest at Barro Colorado I. Not a forest-edge species. Recognition: Unmistakable as Xylopia. With only leaves in hand, it looks just like X. aromatica; bark lenticels and horizontal branches separate it. In habitat and distribution, it resembles X. bocatorena, not X. aromatica.

Xylopia sericea

Xylopia sericea

Xylopia sericea (malagueto). A medium-sized tree

we recently discovered in Panama. Unmistakably Xylopia, with leaves shaped like those of X. frutescens, but with long, white hairs on the underside, horizontal branching, and small buttresses at the base of bigger trees. Distribution: Only known from Cerro Azul and Cerro Jefe and nearby wet lowlands; moreover,

we recently reidentified some trees at Pipeline Rd. as this species (we had identified them as X. frutescens earlier). In Costa Rica known at one site in the north. Appears in forest interior, unlike X. frutescens. Recognition: Leaves very similar to those of X. frutescens, but with prominent white hairs. Xylopia has one other species in Panama, known only in wet forests just east of the Canal; its leaves are much like those of X. macrantha. Six other genera of Annonaceae occur in Panama: Cremastosperma, Klarobelia, Malmea, Porcelia, Sapranthus, and Stenanona. Only the latter two are known in Costa Rica. Except for Porcelia, which is found in dry forest, they are wet-forest or montane species. They are seldom collected in Panama, but Sapranthus viridifolia is quite common in Costa Rica.

Apocynaceae Aspidosperma megalocarpon

Aspidosperma megalocarpon

A large and widespread tropical and temperate family that includes 5000 species of herbs, shrubs, lianas, and many tropical trees. There are 1500 species in tropical America. Familiar among the Apocynaceae are periwinkles (Vinca), oleander (Nerium), milkweed (Asclepias), and the widely planted tropical ornamental frangipani (Plumeria). The family includes 35 species of trees and treelets native to Panama; we cover 16 here, with photos of 12. The single unequivocal character of the family is white latex, which drips copiously from broken parts in nearly all species. Many other families have latex, but only a few produce so much. Leaf traits, though, are not at all clear-cut. As is generally true when

latex is plentiful, leaves tend to be thickened and stiff. All have simple, untoothed leaves, but unfortunately, some species have alternate leaves and others have opposite leaves. This is unusual, because the alternate–opposite dichotomy is usually consistent within plant families. Flowers of the Apocynaceae are reliable, with a characteristic pinwheel form that is familiar in garden periwinkles, that is, each petal emerges from below the petal next to it. Without flowers, and given the variability of leaves, this family illustrates the importance of checking for latex: if there is little or none, your tree is not Apocynaceae; if there is a lot of white latex, Apocynaceae should be your first thought (along with Moraceae, plus the genus Mabea in the Euphorbiaceae). If there is copious white latex and leaves are opposite, it is almost certainly Apocynaceae. But keep in mind Clusiaceae, which always has opposite leaves and yellowish, less-voluminous latex.

Aspidosperma megalocarpon. Not illustrated.

A large forest tree much like A. spruceanum, but with a very distinctive, fluted trunk. Leaves are whitish on the underside. Fruit is somewhat smaller than in A.

Apocynaceae

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Xylopia macrantha

Xylopia sericea

spruceanum. Distribution: Fairly common in wet forests from the Canal Area east. Sparse but widely known in other wet forests. Recognition: Big trees are conspicuous because of their odd trunk, but Minquar-

tia (Erythropalaceae), Platypodium (Fabaceae), and Macrocnemum (Rubiaceae) also have fluted boles. Leaves are like those of A. spruceanum, but much lighter underneath and with fewer secondary veins.

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Apocynaceae

Aspidosperma spruceanum

Aspidosperma spruceanum

Aspidosperma spruceanum (alcarreto, vola-

dor). A large forest tree. Trunk is straight, with light, smooth bark having small lenticels. Leaves are alternate, regularly spaced along branches, very dark green above and much lighter below, with a yellow midvein, and closely spaced, parallel secondary veins that are faint but clear. A good detail is the way secondary veins are alternating: one starts at the midvein but does not reach the leaf margin, the next reaches the leaf margin but not the midvein, etc. Lots of white latex flows from broken leaves, twigs, or fruit. Flowers are tiny, white to yellowish, in dense

terminal heads. Fruit is a large woody capsule filled with winged seeds. Distribution: Common in c. Panama on the Caribbean half, otherwise known from the Osa Peninsula and a few other widely separated locales. A mature forest tree, not in secondary forests or disturbed areas. Saplings are common in the shady forest understory. Recognition: The combination of alternate leaves, faint, closely spaced secondary veins of alternating length, and a yellow midvein make this fairly easy to learn without even checking for latex. Garcinia (Clusiaceae) has remarkably similar leaves, but opposite, not alternate. Ficus (Moraceae) is reminiscent, with yellow midveins and lots of latex, but not at all similar in detail. Three more Aspidosperma species are known from Panama, all rare, and only one is known in Costa Rica. We have not seen any of them, but from specimens it appears their leaves are very similar to those of A. spruceanum, lacking the whitish underside of A. megalocarpon.

Couma macrocarpa

Couma macrocarpa

Couma macrocarpa. Not illustrated. A mediumsized tree. Leaves are large, round, with numerous straight, parallel secondary veins; three leaves emerge together, and the three are equal in size. All broken parts drip latex. Flowers are small, in terminal heads; fruits are hard capsules. Distribution: Only known from a few locations in Caribbean lowlands from the Canal to n. Costa Rica, but fairly common in lowland forests in Bocas del Toro. Recognition: There is nothing else like this, with latex-filled round leaves coming in threes.

Lacmellea panamensis

Lacmellea panamensis

Lacmellea panamensis (leche de vaca, lagarto

negro). A medium-sized forest tree. Trunk is straight and has scattered, somewhat blunt, conical spines. Leaves are simple and opposite, dark green, in a flat plane and spaced regularly along branches. Broken

leaves and fruits produce lots of white latex. Flowers are in clusters on branches at leaf pairs, and have long white tubes. Fruits are berries. Distribution: Widespread in wet and lower montane areas near the Canal and the Osa Peninsula; otherwise sparse in lowland wet sites throughout. Not especially common anywhere, and not in disturbed or secondary forest. Recognition: Opposite leaves arranged regularly in a plane and a spiny trunk are distinctive; the latex eliminates all else but other Apocynaceae. Lacmellea speciosa and the genus Malouetia do not have spines, but both have leaves in a similar arrangement. Leaves of Stemmadenia and Tabernaemontana are also opposite and filled with latex, but are perpendicular to one another and bunched.

Apocynaceae

Aspidosperma spruceanum

Lacmellea panamensis

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Apocynaceae

Lacmellea speciosa

Lacmellea speciosa

Lacmellea speciosa (leche de vaca). Like L. panamensis in most respects, with opposite leaves in a flat plane. But few or no trunk spines, and leaves are larger and have tiny black spots dispersed over the underside. Flowers and fruits as in L. panamensis. Distribution: Sparsely distributed in lower montane forest the length of Panama and Costa Rica, plus a few lowland Caribbean sites; not common. Recognition: See L. panamensis.

Laxoplumeria tessmannii

bling those of the wild cashew, Anacardium exelsum (Anacardiaceae), but producing lots of latex when broken. Flowers are in dense terminal heads. Fruits are very thin, long, cylindrical capsules. Distribution: Only known in wet forests of the Caribbean slope in c. Panama, where widespread but not very common. Can be seen at the outer half of Pipeline Rd., some not far from the road. Recognition: The leaves do not look especially like the ornamental frangipani, and unless you break a leaf to check for latex, you might confuse this with Anacardium excelsum (Anacardiaceae). See also the genus Pouteria (Sapotaceae), all of which have latex and many of which have long, narrow leaves.

Laxoplumeria tessmannii

Laxoplumeria tessmannii (wild frangipani, wild

plumeria). Not illustrated. A medium-sized tree of wet forests. Leaves are long and narrow, densely clustered at the ends of branches, and dark green, resemMacoubea mesoamericana

Macoubea mesoamericana

Macoubea mesoamericana. Not illustrated. A large wet-forest tree. Leaves are thick, wide, oppo-

site, whitish underneath, with prominent, parallel secondary veins. Flowers are white, with short tubes. Fruits are woody capsules with two sections that are broadly connected (unlike the independent sections in Tabernaemontana). Distribution: Known in two sites in the Caribbean lowlands near the Canal, and one in e. Costa Rica. Recognition: The thick, wide, opposite leaves are reminiscent of Couma, but are in pairs, not threes. A second species of Macoubea is known at only a few sites in Caribbean e. Panama.

Malouetia isthmica

Malouetia isthmica

Malouetia isthmica (lechosa). A small or medi-

um-sized forest tree. Leaves are opposite, arranged regularly and in a flat plane along branches. Leaf undersides have tiny pits (domatia) where secondary

veins join the midvein. All parts produce copious latex when broken. Flowers are in dense clusters at each pair of leaves, and have short, white tubes. Fruits are long, thin cylinders. Distribution: Known only on the Caribbean slope of c. Panama in wet and especially swamp forest. Recognition: Opposite leaves in a flat plane, with lots of latex, set Malouetia apart except for Lacmellea, which could be confused. Their fruits are completely different, though. A second Malouetia is known more widely from lowland Caribbean forests throughout Panama and Costa Rica. It is similar to M. isthmica; indeed, we have confused the two species.

Apocynaceae

Lacmellea speciosa

Malouetia isthmica

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Apocynaceae Plumeria rubra

Plumeria rubra

Plumeria rubra (caracucha, frangipani). A familiar ornamental all over the tropics, but native to tropical America and occurring naturally in Panama. A small tree with a bent or leaning trunk. Leaves are long and fairly narrow, thickened, and bunched toward the

ends of the branches (so bunched it is not obvious they are alternate). Leaf veins are yellowish. All broken parts drip white latex. Flowers are familiar, large, fragrant, white pinwheels. Fruits are long, paired woody capsules having winged seeds within. Distribution: Commonly planted in farms and gardens, but can also be seen wild in the driest areas, around Punta Chame or the Azuero Peninsula in Panama and Guanacaste in Costa Rica, where it grows on rocky hillsides. Recognition: Most people get to know this tree in gardens. The leaves from a distance resemble those of Anacardium excelsum (Anacardiaceae), but the resemblance vanishes after feeling the thickness or noting the latex.

Rauvolfia littoralis

Rauvolfia littoralis

Rauvolfia littoralis (lechosa). A medium-sized tree of the dry Pacific coast. Trunk is straight and cylindrical. Leaves are simple, appearing in whorls (four or five leaves emerging together from the branch) and thus may appear palmately compound. Leaves within a cluster are unequal in size. Any bro-

ken part of the tree drips white latex. Flowers are white, in dense terminal heads. Distribution: Occurs widely in forests around Panama City and in c. Costa Rica, and there are a few other records scattered on the Pacific slope. Generally not very common. Recognition: See Couma, which has much wider leaves in whorls of three. No other big tree has whorled leaves. Hamelia patens (Rubiaceae) has clusters of four leaves, but is a bushy treelet and lacks latex. Two other Rauvolfia species are found in Panama. R. purpurascens is a widespread but rarely seen wet-forest species; the other is known only once from c. Panama. All have whorls of four or five unequal leaves, but leaves can be narrow or broad.

Apocynaceae

Plumeria rubra

Rauvolfia littoralis

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Apocynaceae

Stemmadenia grandiflora

Stemmadenia grandiflora

Stemmadenia grandiflora (huevos de gato). A

small tree or shrub of farmland. The stems are small, leaning or bent, and typically forked near the ground. Leaves are small, opposite, with most pairs consisting of one big leaf and one small; adjacent pairs are perpendicular to each other and to the stem. All parts produce lots of latex when broken.

The bright yellow and conspicuous flowers are usually present; sepals below the flowers look like leaves. Fruits are hard, paired capsules. Distribution: Widespread and common in Pacific Panama to s. Costa Rica, especially in dry rural areas; also known on the Caribbean side of the Canal Area. Often in yards and along roadsides but occurs in the forest, although mostly secondary forest. Recognition: Unequal leaf pairs make an excellent trait, because only a few highly divergent species have them. Tabernaemontana arborea has leaves similar in size and shape and also unequal, but it is a big forest tree; the other Tabernaemontana are understory treelets and do not have unequal leaf pairs. Leaves of Terminalia amazonia (Combretaceae) are similar in shape and arrangement to those of Stemmadenia, but should not be confused.

Stemmadenia obovata

Stemmadenia obovata

Stemmadenia obovata (huevos de gato). Simi-

lar to S. grandiflora in most respects. Leaves are somewhat larger and come in unequal pairs. Two leaf traits distinguish this species: a pubescent leaf underside, and secondary veins straight and densely spaced. Distribution: Known only in the driest parts

of Panama, in Herrera and Los Santos. Can be seen along beaches in Los Santos. Also in the dry zone of nw. Costa Rica. Recognition: See the other Stemmadenia, which share the distinctive trait of unequal leaf pairs and lots of latex. Three more Stemmadenia species are found in Panama. S. alfari is a montane species from c. Panama through Costa Rica that has fruits like bull’s horns. S. donnell-smithii is seldom seen in Panama but is widespread in Costa Rican lowlands. The last species is sparsely known from mountain and wet forests throughout. They all have large pinwheel flowers that are yellow or white and opposite leaves with one member of each pair larger than the other.

Apocynaceae

Stemmadenia grandiflora

Stemmadenia obovata

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Apocynaceae

Tabernaemontana arborea

Tabernaemontana arborea

Tabernaemontana arborea (huevos de gato).

A medium-sized forest tree. Trunk is straight, light cream-colored, slightly fluted at the base, with small buttresses. Leaves are opposite, with pairs perpendicular to one another and the branch; often opposite leaves of a pair are dissimilar in size. Secondary

veins are conspicuous, impressed above, curving upward toward the leaf tip. Flowers are small, white pinwheels, easy to find on the forest floor during the reproductive period. Fruits are brown, kidney-shaped, and come in a pair, also easy to find on the forest floor. Distribution: Widespread in the Canal Area and a few wet sites to the east; sparse in lowlands throughout Costa Rica. A tree of mature forest; easy to find at Barro Colorado I. Recognition: The lightcolored and ridged trunk is characteristic and readily learned. The tendency for leaf pairs to be unequal, with lots of white latex, are also easy traits. Stemmadenia are similar in these respects, but are small bushy plants, seldom in the forest. The other Tabernaemontana are small treelets, with leaves quite different from those of T. arborea.

Tabernaemontana undulata

Tabernaemontana panamensis

Tabernaemontana panamensis (huevos de gato). An understory treelet of wet areas. Stem leaning and usually branched at low heights. Leaves opposite, thick, with conspicuous and impressed sec-

ondary veins; pairs are rotated around the branch, but tend to be well-spaced and not grouped toward the end. Lots of latex. Flowers white. Fruits come in pairs of globes (the fruit form that gives several Apocynaceae their Panamanian names). Distribution: Mostly lower montane forests from c. Panama east (Santa Fe, Cerro Campana, and Cerro Jefe); not in Costa Rica. Recognition: Leaves resemble those of the frangipani (Plumeria) in being thick, with conspicuous veins, but not bunched like those of frangipani; does not resemble the other Tabernaemontana and Stemmadenia, but rather Lacmellea and Malouetia. Garcinia (Clusiaceae) has somewhat similar, opposite leaves, but far less latex.

Apocynaceae

Tabernaemontana arborea

Tabernaemontana panamensis

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Apocynaceae

Tabernaemontana undulata

Tabernaemontana undulata

Tabernaemontana undulata (huevos de gato).

An understory treelet of wet areas. Stem leaning and usually branched at low heights. Leaves opposite, narrow and pointed, with yellowish green undersides, arranged in a flat plane and not bunched. All parts drip lots of latex when broken. Flowers can be pink or white. Distribution: Lowland Caribbean wet

forests. Very common at Santa Rita and the upper Chagres River and Bocas del Toro, also the Osa Peninsula. Only in the shady forest interior. Recognition: Opposite leaves in a flat plane are like those of Lacmellea and Malouetia, not the other Tabernaemontana. The color of the undersides is distinctive, not like other Apocynaceae. Mabea occidentalis (Euphorbiaceae) has latex and light leaf undersides, but alternate, not opposite, leaves. Six additional species of Tabernaemontana occur in Panama. All have opposite leaves and fruits in paired capsules. T. longipes is common and widespread throughout Panama and Costa Rica; T. alba and T. amygdalifolia are seen widely in Costa Rica but are very rare in Panama. Those three are wet-forest or montane species. The rest are rarely seen anywhere.

Thevetia ahouai

Thevetia ahouai

Thevetia ahouai (huevos de gato). A bushy tree-

let with bright red fruits. Stem is short, leaning, branched. Leaves are narrow, long, and alternate, thick, dark and shiny above, bunched toward the end of branches and forming dense crowns. All parts have lots of white latex. Flowers are yellow, and the conspicuous fruits can be seen any time of the year. Distribution: Common and widespread in the Canal

Area and east; sparsely known in the west and parts of Costa Rica. Mostly at forest edge, secondary forests, or clearings in mature forest, from dry to wet zones. Often along roads in wooded areas, and easy to find at Pipeline Rd. Recognition: At a glance, the leaves may not be distinctive, but if you remember to check for latex, the narrow, pointed leaves are not difficult to learn. Fruits are obvious and often present. Apocynaceae has two genera of trees that we have not covered: Alstonia and Himantanthus, each with a single species. The former resembles Rauvolfia and is seen most often in mountains of Costa Rica, but there are records from w. Panama; Himantanthus has leaves like those of Aspidosperma but with bold veins; it is very rarely known, only in e. Panama.

Apocynaceae

Tabernaemontana undulata

Thevetia ahouai

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Araliaceae

Araliaceae

Dendropanax arboreus

Dendropanax arboreus

A large and widespread, mostly tropical family that includes trees and shrubs as well as epiphytes, lianas, and herbs. It is the family of ginseng (Aralia) and English ivy (Hedera). The biggest genus, Schefflera, is a common ornamental in gardens around the warmer parts of the world, and is also used as a houseplant. Most Araliaceae are in tropical mountains, and we have only 3 common lowland species in Panama, out of a total of 27 species that qualify as trees or treelets. Many of those 27, however, often (maybe always?) grow as hemiepiphytes—woody trees that grow on other trees. Many Araliaceae have compound leaves, some palmately compound. When not compound, leaves tend to be wide, almost round or rounded at the base, with palmate venation. Leaves or leaflets are often toothed, but some are untoothed. Crushed leaves have a celery-like odor. None have latex. The genus Schefflera is easy to recognize because of its highly distinctive compound leaves.

Dendropanax arboreus (vaquero). A tall forest

tree. Big trees have small buttresses at the base; in many trees, the trunk leans a bit from the vertical. Leaves are alternate, but bunched at the ends of branches. They can be faintly toothed, shiny above, rather thick. Petioles are highly variable in length: short near the end of the branch but much longer on leaves below, thus raising the lower leaves up to the level of the terminal leaves. In small saplings and seedlings, leaves are shaped like mittens, with three lobes. Flowers are small, green to yellowish, in terminal, flat-topped clusters and on stalks of variable lengths. Distribution: Widespread in both Panama and Costa Rica, occurring in virtually every forest, from dry to wet to montane. It is never especially common, and mostly inside the forest, both secondary and old growth. Recognition: The roundish leaves with highly variable petioles are distinctive. The family Hernandiaceae has leaves remarkably like those of D. arboreus in size and shape. There is a small difference at the base of the petiole, and Hernandia has a more limited distribution (but co-occurs with Dendropanax at many sites).

Dendropanax has 12 more species of trees in Panama, plus some small shrubs. The majority are montane species, far less common than D. arboreus, but similar in having petioles of variable lengths.

Schefflera morototoni

Schefflera morototoni

Schefflera morototoni (mangabé). A tall tree of forest edge and clearings. Leaves are unmistakable, palmately compound with 10 large leaflets on long stalks radiating off the end of a branch. Because the entire structure is a single leaf, it makes for one of the largest leaves around outside the palms. The underside of the leaflet is a rich brown. Flowers and berries

are small, but in a big cluster growing above the leaves. Distribution: Widespread on the Pacific slope throughout, and in the Canal Area to the Caribbean. Common along roadsides, open areas, young secondary forests, rare in clearings in mature forest. Abundant in the Canal Area. Recognition: One of the most instantly recognizable trees around. The huge, palmate leaves are highly unusual, and the brown undersides can be spotted on hills far away when breezes turn the leaves over, especially obvious on sunny, dry-season days. Panama has 20 more Schefflera species. Most are epiphytic trees, growing on other trees, from cloud forests, and most species are rare. Leaves are always palmately compound and easy to recognize, but no others have brown leaf undersides like S. morototoni.

Araliaceae

Dendropanax arboreus

Schefflera morototoni

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Arecaceae

Sciadodendron excelsum

Sciadodendron excelsum

Sciadodendron excelsum (jobo de lagarto). A

medium-sized tree of dry and open country. On bigger trees, the bark is thick and furrowed. Leaves are tripinnate, which is very unusual among trees: each leaf is divided into about seven leaflets, each of those subdivided into about seven subleaflets, and each of those divided into five sub-subleaflets. The complete

Arecaceae

leaf is large, up to 2 m long. Leaflets are toothed. Flowers are white or yellowish, in dense, terminal clusters. Berries are round and black. Distribution: Found in dry, open farmland of Herrera and Los Santos in Panama and far nw. Costa Rica, sometimes in fencerows. Also occurs in dry forests, including sites around Panama City. Recognition: Thick, furrowed bark is unusual in the lowland tropics, and there is nothing else with a leaf like this. Araliaceae contains another big genus, Oreopanax, with many species. It resembles Dendropanax, but most are hemiepiphytes. There are occasions where they appear as free-standing trees, for example when growing on rocks, or after the host tree dies (as in strangler figs).

Astrocaryum standleyanum

(Palmae) Astrocaryum standleyanum

palms also differ from all other trees in the way they grow: in most, the trunk does not grow larger in diameter as the tree grows taller. This is helpful in recognizing palms, because it means that the thickness of the trunk is a species character: big palms do not have thin saplings. However, big palms are short when they are juveniles, with the large leaves emerging at ground level.

Astrocaryum standleyanum (chunga, palma Palms are a major tropical forest group, with 2000 species worldwide and 550 in tropical America. Most of the American species are lowland forest trees or treelets, though there is one genus of lianas (Desmoncus; many more are lianas in Asia and Africa). There is no family of tropical trees so thoroughly utilized as palms: their wood is used for building, leaves and stems are used for baskets and thatch, many fruits are edible as is the heart-of-palm, sap is fermented into wine, and the oil palm (Elaeis guianeensis) is one of the tropics’ major crops. Moreover, palms are planted as ornamentals in every city of the tropics and subtropics. Panama has 50 native species of palm trees, of which we describe 11 and illustrate 7. There are many more species introduced from the Old World that can be found along city streets, such a large number we decided we had to omit them here. A good way to learn the ornamentals is to visit Summit Garden near Gamboa. Everyone recognizes palms at a glance, and happily, everyone is nearly always right in so doing. Palms are indeed just what you think they are: unbranched trunks with long compound or fan-shaped leaves only at the top. The few exceptions are still pretty obvious: the spiny liana Desmoncus and the ground palms Geonoma. Even better, palms tend to be quite easy to separate into species, because there are plenty of large and conspicuous distinctions. In addition to the unusual trunk and leaf arrangement,

negra, black palm). A tall, spiny forest palm. The trunk is covered in long, black, very pointed spines, as is the leaf sheath. Leaves are very large, with the typical pinnate, palm form. Individual leaflets are arranged irregularly, with some in clusters of three or four and oriented at various angles from the rachis. Fruits are orange, in large dense clusters just below the leaves. Distribution: Known widely throughout, but not common outside c. Panama. Abundant in forests around the Canal, including old-growth as well as secondary forests near Panama City. Recognition: The only other large native palm with big trunk spines, Cryosophila, has fan-shaped leaves and forked spines. Several Bactris have sharp trunk spines but are much smaller trees. A nonnative Bactris, B. gasipaes, has spines and is nearly as large as A. standleyanum, but it is not in the forest. Leaves of Astrocaryum are the commonest source of fibers for woven baskets in Panama, and the very hard wood is frequently used for building and making spears and bows. Two more Astrocaryum species are found in Panama. A. alatum is common and widespread in the understory of wet Caribbean forests, and can be seen easily at Santa Rita near the Canal or in Bocas del Toro. It has heavy black spines, but is a much smaller tree than A. standleyanum. Another species is known only in e. Panama along the Caribbean slope.

Arecaceae

Sciadodendron excelsum

Astrocaryum standleyanum

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Arecaceae

Attalea butyracea

Attalea butyracea

Attalea butyracea (palma real). The tallest native palm. The thick, gray trunk has slight horizontal constrictions. Leaves are pinnate, very large, with irregularly clustered leaflets. There are no spines anywhere. Fruits are in large clusters just below leaves. Bactris major

Bactris major

Bactris major (uvito, uvito de parra, caña brava,

corozo). Not illustrated. A multistemmed, spiny, forest treelet. Stems are slender, with many long, sharp spines. There are nearly always several stems growing together, connected underground but separate Chamaedorea tepejilote

Chamaedorea tepejilote

Chamaedorea tepejilote (pacaya). A slender

understory treelet. About 3 m tall, the stem is green and divided into sections by horizontal rings. The base of the stem is elevated on small stilts. Leaf is long and has the typical pinnate form of palms. The

Distribution: Mostly a Pacific coast species (the sparse range map is an artifact due to reluctance of botanists to collect the very large leaves and fruits as herbarium specimens). Abundant in secondary forest and farmland in c. Panama; occasional in old growth. Seen throughout the Canal Area, often near roads. Quite common in rural areas, sometimes the only tree remaining in extensive pasture. Recognition: The only tree this can be confused with is the nonnative royal palm (Roystonea regia), which is widely planted in street rows but does not appear outside cities; Roystonea has a gray trunk lacking any horizontal rings. Leaves of Attalea are used to make thatch roofs, which is why it is so often left standing in otherwise treeless farmland.

above. Leaves have the pinnate form typical of palms, and have spines on both the rachis and the stalks of the leaflets. Distribution: Mostly a Pacific slope species throughout Panama and Costa Rica. Widespread in forest understory in c. Panama, across the Canal Area, from dry to wet sites, but not often common. Recognition: Stems growing in clusters, with slender spines, distinguish Bactris. Six more Bactris species are native in forest understory in Panana; three of those reach Costa Rica. Most are smaller than B. major and are not easy to tell apart. A seventh species, B. gasipaes, is not native, but is planted in farmland for its edible fruit; it is much larger but also spiny and multistemmed.

underside of the leaf rachis is green with a yellow line in the center running most of the leaf’s length. Small fruits are on bright orange, branched stalks. Distribution: Throughout Panama in moist, wet, and montane sites, in dense forest shade. Often along streams. Recognition: The green stem and yellow line make this species unmistakable. Many more Chamaedorea species are found in Panama, but most are less than 1.5 m tall and do not enter our tree checklist. We count two others as trees, one in lower mountains, the other in wet forests. All Chamaedorea have slender stems and orange fruit stalks, but few have the yellow line of C. tepejilote.

Cryosophila warscewiczii

Cryosophila warscewiczii

Cryosophila warscewiczii (guágara, palma es-

coba, noli). Not illustrated. A medium-sized, spiny fan palm. Trunk is gray and covered with long spines that are also gray. Each spine can branch into two or more spiny tips. Leaves are fan-shaped. Distribution: Widespread in moist, wet, and lower montane forest. Abundant at a few locations near the Canal, but absent over large distances between them. Recognition: The only native fan palm in lowland for-

Arecaceae

Attalea butyracea

Chamaedorea tepejilote

ests of Panama. The only other tall fan palms are street ornamentals, and they lack spines. Astrocaryum standleyanum has big trunk spines, but they are black and unbranched, and it has pinnate, not fan-shaped leaves. As a juvenile, Cryosophila looks much like the Panama hat plant, Carludovica palmata (which is not a tree and not a palm).

Three other species of Cryosophila occur in Panama. One is known only in Chiriqui and nearby Pacific Costa Rica; one is only from the Darien. The third species has records only in the Canal Area, but we are doubtful it is distinct from C. warscewiczii.

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Arecaceae

Elaeis oleifera

Elaeis oleifera

Elaeis oleifera (corozo, palma aceitera, corozo

colorado, American oil palm). A short, squat forest palm. The trunk is thick, to 30 cm in diameter, and is often prostrate, snaking over the ground for a couple meters. At most, it rises about 2 m above the ground. The leaves have the typical pinnate form of palms and are very long, arching up then hanging back down to the ground. There are thick, short, curved spines along the leaf rachis. Fruits are orange, in a

big cluster at the base of the leaves. Distribution: Mostly a Pacific coast species, though also on the Caribbean coast in c. Panama. Usually in swampy areas in lowland forest. Sometimes seen in pasture where it was presumably left as the forest around it was cleared. Recognition: At a glance, the large leaves near the ground resemble those of a juvenile Attalea butyracea, before the latter has grown a trunk. The curved rachis spines identify Elaeis, though. It also has very regularly spaced leaflets, whereas Attalea and Astrocaryum standleyanum both have irregularly spaced leaflets. Fruits of the oil palm are boiled to produce oil. The African oil palm (Elaeis guianensis) is a relative. It is cultivated in huge plantations in many parts of the world, but we know of none in Panama, though specimens are recorded.

Geonoma deversa

Geonoma cuneata

Geonoma cuneata. Not illustrated. A shrub-sized palm of forest understory. At most 2 m tall and often shorter. The leaves are single long blades, undivided, with two points at the end; they can split or tear and then appear to have leaflets, though. Distribution: Widespread and common in wet and lower montane forest shade throughout Panama and Costa Rica.

Near Pipeline Rd., it grows in extensive groves that have to be pushed through to traverse while hiking. Recognition: Might not look like a palm at a glance, because the leaves can be undivided. But Geonoma does have a woody stem, unlike the other ground plants with similar long leaves (herbaceous species in the Marantaceae, Zingiberaceae, or Heliconiaceae). Many more species of Geonoma are found in Panama, all growing in the same wet-forest habitat, but the majority are less than 2 m tall. We count four species as treelets, all widespread in Costa Rica and Panama. Some have fully divided leaves, like other palms; others have leaves divided just a few times, or divided irregularly.

Iriartea deltoidea

Iriartea deltoidea

Iriartea deltoidea (corneto, pico de negro, pico

de chombo, pene de chombo). A medium-sized stilt palm of wet forests. The trunk is supported on black

stilts that are packed so tightly that no light passes through. Leaves are fairly short given the height of the tree; they are pinnate, with the leaflets arrayed at all angles, producing a brushlike form. Leaflets get wider away from the rachis and are square at the end. Distribution: Wet forests and lower montane sites throughout. Sometimes very common. Recognition: Inside the forest, the two stilt palms, this one and Socratea, will not be confused with any other trees. They are distinguished from each other by the density of the stilts: in Socratea, there is ample space among stilts, but not so in Iriartea. The hard wood of Iriartea is frequently used for building.

Arecaceae

Elaeis oleifera

Iriartea deltoidea

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Arecaceae

Oenocarpus mapora

Oenocarpus mapora

Oenocarpus mapora (maquenqué, trupa). A medium-sized, multistemmed palm. Trunks are 10–15 cm in diameter, gray, and nearly always grow in clusters of up to a dozen (joined below ground but not

above). Leaves are the typical pinnate form of palms. Distribution: Widespread in moist, wet, and montane forest of Panama, including forests near the Pacific around Panama City; reaching far sw. Costa Rica. One of the most common trees in c. Panama: on hilltops near Gamboa it is by far the most numerous tree. Recognition: The medium-sized stems are similar to those of Socratea, but lack stilts. Because they nearly always appear in clumps of many stems, they are easy to identify. Bactris also grows in clumps, but all have spiny trunks. One other Oenocarpus species in Panama is only found in the Darien. It has a single large trunk.

Socratea exorrhiza

Socratea exorrhiza

Socratea exorrhiza (jira, palma de zancos). A medium-sized stilt palm of mature forests. Trunks are gray, 10–15 cm in diameter, with horizontal constrictions spaced regularly, supported on many stilts hav-

ing small, knobby spines. There is ample space among the stilts. Leaves are pinnate, rather short for the height of the tree. Leaflets widen to a square tip. Distribution: Lower montane and lowland wet sites, mostly on the Caribbean slope but also the Osa Peninsula. Very common on the Caribbean half of the Canal Area. Recognition: The two stilt palms (this and Iriartea) are unlike any other species. See Iriartea for the easy and obvious distinction in the stilts. Socratea is the more widespread of the two, reaching further toward the Pacific coast. They are both widespread in Caribbean forests, though, often growing together, and Socratea is usually (but not always) the more common of the two.

Welfia regia

Welfia regia

Welfia regia (conga, palma conga, palma tigre). Not

illustrated. A tall, wet-forest palm. The trunk has alternating horizontal bands of dark and light brown. Leaves are large and long, with the typical pinnate form of palms. Fruits are berries held on woody stalks; the stalks can usually be found on the ground long after the fruits are taken. They are dark brown and have rows of cavities, resembling a large cob of corn after the kernels are gone. Distribution: In wet and lower montane forests; the range map looks misleadingly sparse because the large leaves are difficult to collect. In c. Panama, widespread on the Caribbean half of the isthmus, and fairly common at the outer half of Pipeline Rd. Recognition: Tall and with

very large leaves, similar in size to Attalea, bigger than Astrocaryum. The banding on the trunk is sometimes rather faint and obscured in older trees, but the woody fruit stalks are nearly always present. Welfia is found in mature wet forest, whereas Attalea is most common in secondary forest and open areas on the Pacific coast. They may occur together in forest openings along the Caribbean slope. The remaining palm genera with tree species in Panama (with the number of species in each genus) are Acrocomia (one), Aiphanes (two), Calyptrogyne (one), Colpothrinax (one), Dictyocaryum (one), Euterpe (two), Manicaria (one), Pholidostachys (one), Phytelephas (one), Prestoea (five), Raphia (one), Reinhardtia (one), Sabal (one), Synechanthus (one), and Wettinia (three). Most of those are restricted to Caribbean lowlands; Euterpe can be seen near the Canal on the wet half of the isthmus. Acrocomia and Sabal occur in dry savannas of the Pacific coast, whereas Colpothrinax is restricted to mountain tops. Synechanthus, a skinny little palm with a yellow stem, is seen at Barro Colorado I. and Soberania.

Arecaceae

Oenocarpus mapora

Socratea exorrhiza

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Asteraceae

Asteraceae Piptocoma discolor

Piptocoma discolor

the species are encountered in fields or roadsides, but occasionally in the forest. Fortunately, flowers are often present, making identification fairly easy.

Piptocoma discolor (negro). A small roadside

This is the largest plant family in the world, including daisies, sunflowers, asters, etc. But the vast majority are small herbs, and those that are woody are nearly all small and weak-stemmed. It is an especially important group in temperate or tropical montane areas, but is much less common in lowland rainforests. There are 24 species we call trees in Panama, all weedy species of open areas no more than a few meters in height. Because the group is unimportant in forests of Panama, we cover just one species. The family is best recognized from the composite flowers. Individual flowers are tiny, packed closely together in a head; the sunflower (Helianthus) is a good example. Without flowers, Asteraceae are not easy to characterize. The few tree species tend to have soft, graying leaves that are clustered toward the end of branches. In many species, leaves have irregular lobes, but this is far from universal. Most of

tree of montane areas. Leaves are alternate, densely bunched at the top of the stem. Leaf undersides are silvery gray. Flowers are white, in dense, rounded heads just above the leaves. Distribution: Forest edge and open areas in montane sites, throughout Panama to c. Costa Rica. Recognition: The largest of the Asteraceae in Panama, over 10 m tall at times and looking like a full-fledged tree. Without flowers, it can be learned fairly easily from the very bunched leaves. Another Asteraceae, Verbesina lanata (not illustrated) has similar leaves, but is smaller and has yellow flowers. The other 23 Asteraceae species in Panama that become treelets are in 14 genera (Baccharis, Bartlettina, Calea, Chromolaena, Clibadium, Critonia, Hebeclinium, Koanophyllon, Lasianthaea, Telanthophora, Tessaria, Verbesina, Vernonanthura, and Vernonia). Most are from higher elevations. One, Koanophyllon wetmorei, is a lowland species in the Canal Area, found in small openings in mature forest canopy, but it is rare.

Betulaceae Alnus acuminata

Alnus acuminata

A modest family of trees, almost entirely north temperate, where there are 170 species. Four species occur in mountains of Central American and two of those are also in the Andes. This is the birch and alder family, familiar in North America and Europe, especially far to the north. Panama has just a single species in the alder genus. The family is best characterized by its woody fruits that look like cones of conifers. Leaves are alternate and toothed, and stipules or their scars are

evident. The latter is a useful detail when comparing with several other similar montane trees (Clethra in the Clethraceae and Rhamnus in the Rhamnaceae).

Alnus acuminata (aliso, jaúl). A tall tree of high mountains. Trunk is large, straight, cylindrical, smooth. Leaves are alternate and double-toothed, with one larger tooth where the secondary veins end and a smaller tooth between. The underside is bluish, with prominent yellow, straight, closely spaced secondary veins and straight and parallel tertiary veins. Fruits are woody cones. Distribution: Only in the mountains of Chiriqui through Costa Rica, above 1500 m elevation. Most common in secondary forest, along rivers, or colonizing landslides. Recognition: Will be familiar to anyone who has seen alder in North America or Europe. The veins and teeth are very distinctive. Tall trees with cylindrical trunks high in the mountains are likely this species.

Betulaceae

Piptocoma discolor

Alnus acuminata

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Bignoniaceae

Bignoniaceae Amphitecna latifolia

Amphitecna latifolia

ognize and unlikely to be spotted as Bignoniaceae until you know them well. Flowers of Bignoniaceae are tubular, with the lower lip extended, and usually brightly colored and large. Fruits are often long, thin, and podlike, but are much different in Amphitecna and Crescentia.

Amphitecna latifolia (calabacito, totumillo,

A moderate-sized family, with about 700 species in the world, most in the American tropics. It is rich in lianas but has many trees as well. In the Neotropics, two genera have large trees, Tabebuia and Jacaranda, and both are widely planted as ornamentals. The brilliant purple flowers of Jacaranda are well known in cities of the tropics and subtropics. In North America, the family also is known for the catalpa tree (genus Catalpa) and the trumpet creeper vine (Campsis). Twenty-four native species of bignons are trees in Panama, of which we cover 7, and we add a description of a widely planted ornamental from Africa. Most Bignoniaceae have opposite, compound leaves, which is an unusual configuration among trees. Only a few other species share this trait (Staphyleaceae and a few Fabaceae in lowland forests), so Bignoniaceae are readily recognized. But to keep matters interesting, there are some trees with simple, alternate leaves in this family, and without flowers, Amphitecna and one Tabebuia are difficult to rec-

maraquita de marea). A small tree with a trunk that forks near the ground. Leaves are alternate, simple, dark green above, broadest near the tip but with a tiny point. Flowers are large, greenish, held on larger branches. Fruit is a large, woody gourd. Distribution: Along both coasts, where it occurs near mangroves or beaches, sometimes in farmland. Occasionally in wet lowland forest away from the coast. Recognition: This species is difficult to recognize, because it certainly does not recall the typical leaf pattern of the familiar Bignoniaceae. Fortunately, it often has fruits, and they are distinctive. See Crescentia, which has a similar fruit. Three other Amphitecna species are found in Panama. A. spathicalyx is similar in most respects, though with a larger and more elliptical fruit; it is mostly a lower montane species, found only in c. Panama. The other two species are widespread lower montane and wet-forest species, in both Panama and Costa Rica.

Crescentia cujete

Crescentia cujete

Crescentia cujete (calabazo, totumo). A straggly

treelet of farmland with a well-known fruit. Leaves are long and narrow, much broader toward the tip, thick and shiny above, simple and alternate. Flowers

are large, greenish, on stems below the leaves. Fruits are large, woody globes used for making bowls. Distribution: Widespread around human habitation, often cultivated, native to the American tropics but its natural origin obscure. Not in the forest. Recognition: See Amphitecna, but the two relatives are quite different. From a distance in its agricultural habitat, Crescentia usually can be spotted by the multiple, straggly stems. Fruits are often present. A second species of Crescentia, C. alata, is common in dry nw. Costa Rica but rarely seen in Panama. It also has big, woody fruits, but its leaves are trifoliate, with winged petioles.

Bignoniaceae

Amphitecna latifolia

Crescentia cujete

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Bignoniaceae

Jacaranda copaia

Jacaranda copaia

Jacaranda copaia (jacaranda, pata de elefante,

palo de buba, nazareno, guabanday). A tall forest tree with spectacular purple flowers. The trunk has yellowish bark with narrow, vertical fissures. The base is expanded into little buttresses, and these have horizontal wrinkles resembling an elephant’s trunk. Leaves are large, opposite, bipinnately compound, that is, the whole leaf is divided into leaflets, and the leaflets are divided into subleaflets, the latter tiny but the entire leaf up to 1 m long. Flowers in March and April for a few weeks, producing dense displays of deep purple visible for kilometers (easily visible, for

instance, in satellite images and aerial photographs). Distribution: Very common in moist and wet forest of c. Panama. Can be abundant there as saplings along roadsides and clearings in wet or moist areas. Elsewhere, sparsely but widely known. Recognition: The big, bipinnate leaves are easy to spot. Check to make sure the leaves are opposite, because Schizolobium parahyba (Fabaceae—Caesalpinioideae) has similar, alternate leaves (the two species are often confused as saplings). Inside the forest, the bark and elephant-trunk base are easy to learn. During the dry season, purple flowers pile up beneath crowns, sometimes carted away in purple columns by leafcutter ants. Another species of Jacaranda, J. caucana, is planted as an ornamental in Panama, commonly in the Canal Area. It is South American, but may grow naturally in forests of Darien, Chiriqui, and sw. Costa Rica. It has bipinnate leaves, but the subleaflets are tiny. J. mimosifolia is also sometimes seen around pastures or yards, but is native only to South America.

Parmentiera cereifera

Parmentiera cereifera

Three other species of Parmentiera are found in Panama, and two reach Costa Rica; all are scarce, but P. macrophylla occurs at Santa Rita. All have trifoliate leaves, but with larger leaflets than P. cereifera, and all have odd, long fruit pods.

Spathodea campanulata (African tulip tree).

Parmentiera cereifera (árbol del vela). A small

forest tree with a trunk forked near the ground. Leaves are opposite, compound, with three leaflets; the petiole is winged. Flowers are large, white, appear on the trunk and large branches, sometimes near the ground. Fruits are bizarre: yellow, long, thin, waxy, and often present. Distribution: Only known from the Canal Area, where it is fairly common on limestone outcrops near Madden Dam and Fort San Lorenzo. There are a few trees in the forest around Gamboa. Recognition: Fruits are unmistakable when present. Even without fruits, opposite, trifoliate leaves are very unusual.

Not illustrated. An alien tree, native to Africa, planted here as an ornamental. Has a straight trunk and opposite compound leaves with many leaflets in an odd-pinnate arrangement. At the base of each leaf, there is a round, miniature leaf, which is actually the stipule. Large, bright orange flowers are in clusters above leaves. Fruits are long pods that stick upward out of the crown. Distribution: Widely planted throughout the tropics and subtropics, and seen commonly in cities and towns. Recognition: The leaves are quite similar to those of Spondias mombin (Anacardiaceae), which also grows commonly along streets of the Canal Area, but leaves of Spathodea are opposite, unlike Spondias and several other trees of the area with similar compound leaves. Other species with opposite, compound leaves are different enough so as not to be confused with Spathodea.

Bignoniaceae

Jacaranda copaia

Parmentiera cereifera

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Bignoniaceae

Tabebuia guayacan

Tabebuia guayacan

Tabebuia guayacan (guayacan). A huge forest

tree with an unparalleled floral display. The straight trunk can be more than 1 m in diameter and trees may be 40 m tall. The wood is incredibly hard, like stone, and dead trees remain standing for decades, including some in Gatun Lake, killed by rising waters 100 years ago. Bark is brown, furrowed. Leaves are palmately compound, with five leaflets. Flowers are bright yellow, produced after rains during the dry season, from early March into May; many trees flower synchronously, then drop all their flowers after 1 or 2

days. Trees are leafless when they flower (or sometimes a large leafless branch will flower while another branch on the same tree has new leaves but no flowers). Crowns are completely covered in yellow when flowering, and easily visible in satellite images. The following day, the ground below is a yellow carpet. Flowers do not have nectar, but fool pollinators because of their immense and sudden appearance. Distribution: Very common around the Canal and less so westward on Panama’s dry Pacific slope. Sparse elsewhere and seldom seen in Costa Rica. It is a common street tree in Panama City. Recognition: The leaves are readily recognizable, even in tall trees, but note that other Tabebuia are very similar. This species always flowers in March and later, and nothing else produces such brilliant yellow displays at that time; in December, Schizolobium parahyba (Fabaceae—Caesalpinioideae) produces large displays of yellow flowers, and is often confused, but the two flowering periods do not overlap.

Tabebuia ochracea

Tabebuia ochracea

Tabebuia ochracea (guayacan). A large tree of

the dry Pacific slope. Very similar to T. guayacan, but

not as large. Leaflets can be faintly toothed, and petioles always have fine reddish hairs, unlike T. guayacan. The yellow flowers have red lines on the lower petals, unlike T. guayacan. Distribution: Mostly in the dry zone of c. Panama and nw. Costa Rica, in pastures and dry forests; there are some trees around Panama City. Recognition: If you are not paying attention, you will usually think this is T. guayacan; you have to look closely at leaves or flowers to be sure, although in dry pasturelands, it will be T. ochracea and in forests, T. guayacan. See also Tabebuia rosea.

Bignoniaceae

Tabebuia guayacan

Tabebuia ochracea

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Bixaceae

Tabebuia rosea

Tabebuia rosea

Tabebuia rosea (roble). A large forest tree with pink flowers. Bark is grayish, furrowed. Leaves are palmately compound with five leaflets, typical for the genus. Flowers are shaped just like those of the other Tabebuia, but pink and sometimes almost white. Quite unlike T. guayacan, this species produces a few pink flowers for many weeks during the dry season, in small numbers at any one moment; it is leafless when flowering. The wood is hard, but unlike the guayacan, is widely used for timber. Distribution: Occurs nearly everywhere in Panama and Costa Rica, in farmland and forest, wet, moist, and dry. Not especially common inside the forest. Very often planted along streets and in gardens, farms, and plantations, much more so than guayacan or jacaranda. Recognition: Throughout the dry season, the pink flowers will become familiar, because this tree is everywhere. Without flowers, more difficult to

identify, because the leaves are similar to those of the guayacan. The bark of the guayacan is brown and furrowed; this species gray and furrowed; guayacan leaves are a bit more papery and lighter colored than those of the roble. Because the trees are seen so often, you will learn to tell them apart. Four more Tabebuia species are known in Panama. Most have opposite, compound leaves with five leaflets (though sometimes three to seven). T. striata, however, has simple leaves and without its white, tubular flowers will not be recognized to genus or family. It is found only at Santa Rita and east along the Caribbean slope; the other species are rare in Panama but seen more often in Costa Rica. Two more native tree genera in the Bignoniaceae are found in Panama, Godmania and Tecoma, both with a single species. T. stans is a well-known treelet, often in gardens and farms, with yellow, trumpetcreeper flowers similar to the guayacan, but with pinnate leaves having three to five leaflets. Godmania aesculifolia is a tree of the driest zones of both countries, with leaves like those of the guayacan. Finally, Kigelia africana is an alien from Africa with brown, sausage-shaped fruits dangling from long brown threadlike stalks; it is occasionally seen along city streets.

Bixaceae

Bixa orellana

Bixa orellana

A very small, tropical family of treelets or shrubs, having fewer than 10 species, of which 8 or so are from the Americas. There is just one genus with two species in Panama. It is sometimes merged with Cochlospermaceae. We will not review family traits in such a small family.

Bixa orellana (achiote). A small shrub of gardens and farms. The small, leaning stem is multiply

branched near the base. Leaves are simple, alternate, heart-shaped, with three to five main veins radiating from the base. When broken, orange sap flows out. The fruit is a dry, spiny capsule. Inside are small seeds that produce a thick, red, paintlike oil when squeezed. Distribution: Lowlands throughout, mostly around human habitation, in gardens and farms, not in the forest. Recognition: The big, heartshaped leaves with conspicuous veins on a weak, multibranched shrub are easy to learn. Fruits are often present and distinctive. The other species of Bixa, B. urucurana, has similar leaves but is a somewhat larger treelet and occurs in the forest, widely distributed though not commonly found.

Bixaceae

Tabebuia rosea

Bixa orellana

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Boraginaceae

Boraginaceae Bourreria costaricensis

Bourreria costaricensis

A large, worldwide family, mostly herbs but with a few genera of trees, shrubs, and lianas, and mostly tropical but with a few temperate herbs. There are 27 species of trees and shrubs in the Panama checklist, 22 of which are in the genus Cordia. We cover nine Cordia species and one Bourreria. As a family, Boraginaceae are rather diverse and difficult to describe. Considering only trees, it is most useful to focus on the big genus Cordia. They all have simple, alternate leaves, sometimes toothed. Many of the Cordia have a conspicuous and unusual leaf pattern, where one leaf emerges at the fork of two branches (a leaf at each branch dichotomy). Also, many Cordia have several branches emerging together from the trunk (verticillate branching), with a swollen node at the point of emergence; some species have ants inhabiting those nodes. Moreover,

most Boraginaceae tend to have scaly leaves, rough to the touch. None of these traits, however, is universal in Cordia, and the last two are common in other families. However, a leaf arising from a branch dichotomy is a sure bet for Cordia. Most Boraginaceae produce flowers in a characteristic pattern called a cyme, but Cordia does not, so if you are studying trees, you do not need to learn what a cyme is.

Bourreria costaricensis (canalú). A medium-

sized tree of wet Caribbean forests. Leaves are alternate, rounded at the tip, smooth and shiny, with few secondary veins. The trunk is ribbed, with canals running vertically. Flowers are large and white. Distribution: Widespread but not especially common in Caribbean wet forests throughout Panama and especially in nw. Costa Rica; not seen on Pacific slope. Recognition: The ribbed trunk is the best trait; see Aspidosperma megalocarpon (Apocynaceae) of wet forests with a similar trunk. Leaves of Bourreria are not distinctive, resembling some Euphorbiaceae or Drypetes (Putranjivaceae). Bourreria oxyphylla also occurs in Panama, and is known only from far e. Panama and far n. Costa Rica. Flowers are similar to those of B. costaricensis, and it also has a ribbed trunk.

Cordia alliodora

Cordia alliodora

Cordia alliodora (laurel). A well-known native tree, often in plantations. The trunk is smooth and light colored, never quite straight, but unbranched until the upper parts. Smaller plants are characterized by swellings along the stems where the branches exit; three or four branches exit from a single swelling. Ants inhabit the swollen stem node. Branches are not straight where they exit, but rather arch upward. In larger trees, this same pattern persists, but is not always clear. Crowns of larger trees are small, rounded, with irregular and arching branches. Leaves are oval, alternate, bunched, and leaves and petioles are scaly and rough to the touch, with small hairs. Flowers are produced in large masses above the leaves in February. They are conspicuously bright

white at this time, but fade to a dull brownish while remaining on the tree for weeks. Individual flowers are small, white, with green stripes. Foliage in the crown becomes very thin during April and May, and the trees are completely leafless for a period until July. Distribution: A common and widespread edge and roadside species, especially on the Pacific slope, throughout Panama; also fairly common in natural canopy openings in mature forest. Occasionally on the Caribbean side, too. Very common along roads near Panama City. Recognition: Takes awhile to learn well. The leaves are not at all distinctive. Most important in smaller plants is the branching pattern and the swollen stem nodes. Larger trees are easily recognized during February and March, when they are covered in dense white flower clusters. Later, when the crown is leafless, it can be recognized, because no other common species are leafless at the same time, and by the small, dense crown of twisting branches. When there are leaves and no flowers, it takes a good deal of experience to learn to recognize it as an adult (when the swollen stem nodes are not evident). The wood is good quality and frequently used in various kinds of construction. It is grown in plantations in both Central and South America.

Boraginaceae

Bourreria costaricensis

Cordia alliodora

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Boraginaceae Cordia bicolor

Cordia bicolor

Cordia bicolor (muñeco). A medium-sized tree of

leaf emerges from each branch fork. Leaves are rough to the touch. Flowers are very small, white, in dense terminal clusters. Fruits are small berries. Distribution: Very common around the Canal from the middle to the Caribbean side. From c. Panama through Costa Rica, it is widespread in lowlands of both slopes, but sparse. Recognition: At a glance, leaves are similar to those of C. lasiocalyx, which is also common around the Canal. But leaves of C. bicolor lack the long, pointed tip, and leaves of C. lasiocalyx are smooth.

wet and moist forests. On many branches, a single Cordia cymosa

Cordia cymosa

Cordia cymosa (laurel negro). A medium-sized

tree of wet and lower montane forests. Branches are

verticillate, with three or four emerging from the trunk at one spot. Leaves are broad, sometimes nearly round, rough to the touch, with parallel but widely spaced secondary veins; in juveniles, leaves sometimes have teeth. One leaf routinely emerges at the fork of two branches. Small white flowers resemble those of other Cordia. Distribution: A lower montane species of c. and w. Panama through Costa Rica, but also in wet lowlands. Fairly common in cloud forest at Cerro Campana. Recognition: As Cordia by the characteristic leaf at branch nodes; this species has wide, rounded leaves.

Boraginaceae

Cordia bicolor

Cordia cymosa

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Boraginaceae Cordia dentata

Cordia dentata

Cordia dentata (biyuyo). A small tree of the dry

parts of Panama. Does not resemble other Cordia ex-

cept for the asperous leaves; does not have verticillate branching or leaves at branch forks. Instead, recognized by its smallish leaves with three or four widely spaced secondary veins. Leaves are toothed, but teeth are sparse and not especially conspicuous, sometimes only toward the leaf apex. Yellow flowers are in small terminal clusters. Distribution: Occurs in the driest parts of Panama and Costa Rica, along roads, fencerows. Recognition: Lacking the typical characteristics of the genus means this is difficult to recognize as Cordia. In its narrow habitat, few other species have toothed leaves.

Cordia dwyeri

Cordia dwyeri

Cordia dwyeri (laurel negro). A medium-sized tree of wet and lower montane forests. Much like C.

cymosa in trunk and branching, including asperous leaves at branch forks. But leaves are quite different in shape, being narrow with a pointed tip, blunt teeth, and veins impressed above. Leaf underside has a golden tint from dense hairs. Flowers are small and white, like those of the other Cordia. Distribution: Known sparsely from lowland Caribbean and some lower montane sites. Recognition: Has the characteristic leaf at branch nodes like several congeners, but narrow leaves with depressed veins and a golden underside distinguish this species.

Boraginaceae

Cordia dentata

Cordia dwyeri

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Boraginaceae Cordia eriostigma

Cordia eriostigma

Cordia eriostigma (paico). A small tree of lower montane forests. Leaves are pointed, small, and ap-

pear at branch forks as in many other Cordia, but are not rough. They have few secondary veins that arch well forward. Flowers are white and just like those of other Cordia. Distribution: Wet and montane forest in c. and w. Panama, plus c. Costa Rica; fairly common in the cloud forest at Cerro Campana. Recognition: The leaves have the distinctive Cordia pattern, appearing where branches fork. Leaves of Cordia lasiocalyx are similar, but that is a lowland species. Leaves of other montane and wet-forest Cordia (C. cymosa and C. dwyeri) also appear in branch forks, but have different venation.

Cordia lasiocalyx

Cordia lasiocalyx

Cordia lasiocalyx (paico). A small tree of lowland

moist forests. Leaves are typical for Cordia in most respects, with a prominent leaf at branch forks, a few

secondary veins arching forward, thin and papery, pointed at the tip. They are not rough to the touch, like many congeners. Branchlets holding leaves are green. Flowers and fruits are white. Distribution: Common around the Canal from Barro Colorado I. to Panama City, plus at a few lower montane sites in e. Panama. Only one record from far n. Costa Rica. Recognition: Quickly recognizable as Cordia from the leaf arrangement. It is most like C. bicolor, also of lowland moist forests and with similar, but rough leaves. See also the leaves of the montane C. eriostigma, but the latter has red fruits.

Boraginaceae

Cordia eriostigma

Cordia lasiocalyx

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Boraginaceae

Cordia panamensis

Cordia panamensis

Cordia panamensis (niguita, muñeco, lengua de

vaca). A small tree of the dry zone. Typical for Cordia,

branches are verticillate, and leaves are rough to the touch, and some leaves emerge at branch forks. But leaves are quite a bit larger than those of other species in the genus, with many parallel secondary veins that barely arch upward. Small white flowers in big clusters are typical Cordia; fruits are white berries. Distribution: Widespread in Pacific coast dry forests, often along roads; quite common around Panama City. Also in the dry zone of nw. Costa Rica, and on the Caribbean slope near the Canal. Recognition: The leaf arrangement quickly identifies this as Cordia; the large leaves are distinctive for the genus.

Cordia porcata

Cordia porcata

Cordia porcata (paico). A treelet of wet Caribbean forests. Leaf and branch patterns are typical for Cordia, but leaves are smooth, shiny, and lack rough hairs; they are highly variable in size. Flowers are white, in small clusters. Distribution: In Caribbean wet forests or lower montane sites in c. Panama and throughout Costa Rica. Recognition: C. lasiocalyx is the other Cordia with smooth leaves, but in C. porcata, leaves are by comparison narrow, smooth, and

shiny. Moreover, C. lasiocalyx is not typically in really wet forests. We just covered nine Cordia species. There are another 13 species in Panama, with 9 of those reaching Costa Rica. Three species are fairly widespread in both countries: C. collococca in Pacific lowlands, C. lucidula in wet forest, and C. spinescens in the mountains. Other species are rarely seen. Some are readily recognized from the leaves at branch forks, but not all, and as should be clear from the nine species we illustrated, leaf form and pattern are quite variable across the genus, making it a difficult group to get to know. Two more genera of Boraginaceae are considered trees, Ehretia with one species and Tournefortia with two. Ehretia is a good-sized tree with very rough leaves, known mostly in Costa Rica but just reaching w. Panama. T. glabra is a small, cloud-forest tree with white berries that is found widely. The other Tournefortia is known only from a couple sites in Panama.

Boraginaceae

Cordia panamensis

Cordia porcata

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Burseraceae

Burseraceae Bursera simaruba

Bursera simaruba

A moderate-sized, almost entirely tropical family, with 650 species worldwide and 300 in tropical America. Nearly all are trees, and they are an important component of most tropical forests everywhere and widespread in Panama and Costa Rica, with several species very common. There are 14 species native to Panama; we cover 6 here. This is one of easiest families to learn, because it is very consistent in a few key traits. Leaves are always compound, with few leaflets (generally no more than nine), and they are odd-pinnate, meaning there is a terminal leaflet. Whereas leaves are alternate, leaflets are always opposite. Leaves, young twigs, and fruits have a turpentine-like odor (as in Anacardiaceae), though in some species, and especially in older leaves, it can be faint. In the most

common rainforest genera, there is a swollen node on the leaf rachis where the last pair of leaflets are held, visible from the underside. Flowers are small, held in terminal clusters, and not distinctive. Fruits are capsules that split open to reveal seeds carrying a pulplike aril; the fruit capsule usually has a stronger odor than leaves.

Bursera simaruba (indio desnudo, almácigo, gumbo limbo). A conspicuous and very well-known tree of drier forests. Bark is smooth, red, peeling in sheets; the inner bark beneath is shiny green. Leaves are compound, with five to nine leaflets having long, pointed tips. Distribution: Common and sometimes abundant in the dry Pacific section of c. Panama and nw. Costa Rica. Very common around Panama City, and often planted in fencerows in rural areas. Also occurs on the Caribbean slope on limestone or rocky soils. Recognition: The bark will never be confused. The related Bursera tomentosa also has smooth peeling bark, but gray with only a hint of red. It is only found in the very driest areas, and also appears in fencerows there. A third species, B. inversa, is known only in the Darien.

Protium panamense

Protium costaricense

Protium costaricense (copal, chutra, kerosín, alcanfor). A medium-sized forest tree of wet and moist forests. In older trees, the base of the trunk has stiltroots. Leaves are alternate, odd-compound, and have swollen nodes where leaflets meet the leaf rachis.

Leaflets are small compared to the next two Protium. Young twigs have a thin mat of fine red hairs. Distribution: Widely known in lowlands of Costa Rica, but in Panama largely restricted to Barro Colorado I. and nearby Soberania. In forest interior only. Recognition: Species of the genus Protium are easy to recognize from the compound leaves and the swollen nodes along the leaf rachis, where each leaflet pair meets. Within the genus, species can be difficult to recognize. This species is quite similar to P. glabrum and P. confusum (not illustrated); the red hairs on young twigs are distinctive, but not helpful on older twigs. P. panamense and P. tenuifolium have much larger leaflets.

Burseraceae

Bursera simaruba

Protium costaricense

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Burseraceae

Protium panamense

Protium panamense

Protium panamense (copal, chutra). A medium-

conspicuous stilt-roots. Leaves are alternate, odd-pinnate, and have swollen nodes where leaflets meet the leaf rachis. Leaflets are large. The tips of the leaflets have only a short point. Distribution: Widespread on the Caribbean half of the isthmus, plus the wet Pacific slope of sw. Costa Rica. Abundant at Barro Colorado I. and elsewhere in the wet half of the Canal Area. In forest interior only. Recognition: Again, easy to recognize as Protium. This and P. tenuifolium are very similar, but the latter lacks stilt-roots and has a long narrow tip on each leaflet.

sized tree of wet and moist forests. Older trees have Protium tenuifolium

Protium tenuifolium

Protium tenuifolium (copal, chutra). A mediumsized forest tree of wet and moist forests. Unlike other Protium, never has stilt-roots. Leaves are alternate, odd-pinnate, and have swollen nodes where leaflets meet the leaf rachis. Leaflets are large, and the tips of the leaflets are greatly extended. Distribution: Very common around the Canal and east; only sporadic

records from Costa Rica. Seen widely around Panama City but not other dry sites; also known at some lower montane and wet sites. Recognition: See the very similar P. panamense. P. tenuifolium lacks stiltroots, but this is no help in small plants, which never have stilt-roots in any species. Panama has 13 other Protium species, all with five to seven leaflets on odd-compound leaves. P. glabrum and P. confusum are lowland wet-forest species quite common on the Caribbean side of the Canal Area and throughout Costa Rica. They are similar to P. costaricense but lack twig hairs. We also recently identified P. trifoliolatum in wet forests east of the Canal, a South American species with only three leaflets previously unknown in Panama.

Burseraceae

Protium panamense

Protium tenuifolium

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Burseraceae

Tetragastris panamensis

Tetragastris panamensis

Tetragastris panamensis (animé, cuatro estomagos, chutra, kerosín). A dominant old-growth canopy tree. The trunk is tall and straight, with gray, slightly furrowed bark and modest plank buttresses at the base. Leaves are alternate, odd-pinnate, generally with 7–11 opposite and fairly small and narrow

leaflets. Like Protium, there are swollen nodes where leaflet stalks meet the leaf rachis. Distribution: A very widespread species in Panama and Costa Rica, in all kinds of forest from dry and wet to lower montane. At Barro Colorado I., it is one of the dominant trees in the old-forest canopy, and saplings are abundant in understory. It is also very common near Panama City and on islands off the Pacific coast, but shows up all over in lowland wet forest of the Caribbean slope too. Recognition: Leaves are just like those of Protium, but leaflets are smaller and narrower. No Protium reaches such a large size. Because of its abundance, big trees are not difficult to learn from the gray bark and small buttresses. Also look for the fallen green fruit capsules, which are fragrant when crushed.

Trattinnickia aspera

Trattinnickia aspera

Trattinnickia aspera (caraño). A large canopy

tree of moist and wet forests. Trunk is straight and tall, with small buttresses. During the rainy season, the bark exudes a fragrant, black resin that accumulates on the trunk at the base. Leaves are alternate, odd-pinnate, with many long, narrow leaflets that are

hairy and rough to the touch. The leaflet stalk has a groove on the upper side. Distribution: A wet-forest species but only known near the Canal, in Bocas del Toro, and the Osa Peninsula. Common at Barro Colorado I. and Soberania. Recognition: The leaves are rather different from those of other Burseraceae, with more leaflets and rough hairs. More likely to be confused with Mosquitoxylum jamaicense (Anacardiaceae), which has many hairy leaflets; Trattinnickia may also look like Spondias (Anacardiaceae), but the latter has smooth leaflets with no hairs. Trattinnickia burserifolia is an Amazonian species that we found at one site at Santa Rita, and there is one more record in the mountains of Panama.

Burseraceae

Tetragastris panamensis

Trattinnickia aspera

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Cannabaceae

Cannabaceae Celtis schippii

Celtis schippii

A small family of about 100 species worldwide, nearly all trees or shrubs, but also a few lianas. They are mostly Asian; in the American tropics, there are just 12 species. This is the hemp family, including marijuana (Cannabis sativa) and also hops (Humulus). The elm family, Ulmaceae, is closely related, and the Neotropical Cannabaceae were all once classified with the elms. There are five species of Cannabaceae in Panama that are trees, and we cover two. The Neotropical species all have simple leaves with three major veins arising from the base. Some have alternate leaves, but one has opposite, and some but not all have teeth. Flowers are tiny.

Celtis schippii. A medium-sized canopy tree. Leaves are simple, alternate, without teeth, and have three main veins arising from the base, two of them arching forward toward the leaf tip. The tertiary veins are ladderlike, crossing between the three main veins or to the major secondary vein. Flowers are solitary or in pairs at leaf bases, with white petals. Distribution: Mainly known in c. Panama, but there are a few records from wet forests of Costa Rica. In forests around Barro Colorado I. and Soberania it is usually in the understory and not very common. Recognition: The venation is distinctive. See several Euphorbiaceae (Acalypha and Alchornea) and Salicaceae (Hasseltia) with similar leaves but generally toothed; for example, Hasseltia looks just like a toothed Celtis. The Malvaceae all have three major leaf veins, and Quararibea or Theobroma might be confused with Celtis. See also Muntingia (Muntingiaceae). Melastomataceae have the same style venation, but much denser ladderlike tertiary veins. Another Celtis species is typically a liana, but sometimes grows as a small, weak-stemmed tree. It has spines and toothed leaves.

Trema micrantha

Trema micrantha

Trema micrantha (jordancillo, capulín). A tall roadside tree. Trunk is straight and generally branched only at the top. The crown is thin, with much light passing through. Leaves are simple, alternate, and spaced regularly in a horizontal plane along the branches. They are toothed, and have three prominent veins arising from the base. The leaf base can be asymmetric, but not always. Flowers are white, tiny; fruits are berries. Distribution: Everywhere in Panama and Costa Rica, from the driest to the wettest plus lower montane. Common along

wooded roads, open areas, and forest edge. Inside the forest, only found in large canopy openings, never in the shade. Recognition: The toothed leaves with three main veins are very good characters, but Trichospermum galeottii (Malvaceae—Grewioideae) is remarkably similar and grows in the same habitat. The biggest difference is that leaves of Trema are widest near the base, whereas leaves of Trichospermum are widest toward the tip; also, Trichospermum has conspicuous purple flowers. See also Muntingia (Muntingiaceae). Trema is one of the fastest growing trees in the world. A second species of Trema is very similar. We recently reidentified trees at Barro Colorado I. as this other species. In addition, one more genus of Cannabaceae is native to Panama, Lozanella, with a single montane species with toothed leaves much like those of Trema.

Cannabaceae

Celtis schippii

Trema micrantha

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Capparaceae

Capparaceae Capparis cynophallophora

(Formerly Capparidaceae)

Capparis cynophallophora stamens. Capers are related to the mustard family, and an odor of mustard oil might be detected. Without flowers, any treelet with variable petiole lengths or colorful scaling under the leaves should bring Capparis to mind. The species with leaves in a regular flat plane lacking scales, however, look like many other families.

The caper family includes 600 species worldwide, with 145 in tropical America. Most are shrubs of dry forests or desert, with only a few rainforest species; some become tree-sized, and a few are herbs or lianas. Panama has 21 species of trees and shrubs, and we cover 5. Capparaceae are very difficult to characterize as a group. Crateva has compound, trifoliate leaves, but Capparis has simple leaves. About half the Capparis species have petioles that vary greatly in length, an excellent character; however, it holds for only one of the species we cover here. Leaves tend to be thick and in about half the species are scaly below, sometimes with a silvery or golden color. Flowers are large and striking, with elongated

Capparis cynophallophora (carne de venado). A treelet of dry areas. The small, often forked trunk can be slightly ribbed, and so can branches. Leaves are shiny green above, whitish, brown, or golden below due to rough scales. Secondary veins are inconspicuous or invisible (especially below). Branchlets have small, rough scales. Flowers and fruits are striking and conspicuous when present; look for the long fruit pods with constrictions between seeds. Distribution: Mainly grows in the driest Pacific zone, and also appears at coastal sites and on rocky soils where moisture is scarce, even in higher-rainfall areas. Recognition: The leaves are easy to recognize; a few other species have golden-tinted undersides (Cordia dwyeri in the Boraginaceae and Chrysophyllum cainito in the Sapotaceae), but not with such faint secondary veins. The distribution in very dry areas is a key trait.

Capparis frondosa

Capparis frondosa

Capparis frondosa (garrotillo). A treelet of moist-

and dry-forest understory. Trunk is short and forked. Leaves are bunched at the end of branchlets, and those near the branch tip are smaller and have shorter petioles than leaves below, so that lower leaves are elevated to the same height as the upper ones. The

petiole is swollen at both ends. Flowers and fruits are similar to those of C. cynophallophora, but smaller, and the fruit is much less elongated. Distribution: Grows in forest shade in dry regions along the Pacific slope in Panama and Costa Rica. Also abundant at Barro Colorado I. and appears in wetter areas where soils are rocky (for instance, at Fort Sherman near the Caribbean). Recognition: Leaves varying greatly in size and petioles varying greatly in length are unusual and very distinctive. Some other Capparis share this trait, but none we are treating here. The common and widespread tree Dendropanax arboreus (Araliaceae), which is not related, has the same leaf arrangement, and will be confused if you are not careful. But leaves of Dendropanax are thick, rounded, and shiny, not like those of Capparis.

Capparaceae

Capparis cynophallophora

Capparis frondosa

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Capparaceae Capparis indica

Capparis indica

Capparis indica (carne de venado). A treelet of dry regions. Similar to C. cynophallophora, but the scales on leaf undersides do not have a golden tint, but are rather greenish or sometimes silvery. Petioles

are slightly swollen and have a channel on the upper side, but only slightly different from C. cynophallophora. Flowers are white, seed pods long, thin, and constricted. Distribution: Nearly entirely restricted to dry Pacific coast sites in Costa Rica and Panama. Usually in forest. Recognition: If recognized as Capparis, it makes a nice comparison with the similar, but golden-leaved, C. cynophallophora. The difficulty will probably be recognizing the genus, though: without flowers or fruits, the alternate leaves in a horizontal plane are not distinctive. You have to notice the rough, scaly leaf undersides, present in many Capparis but in few other species.

Capparis pittieri

Capparis pittieri

Capparis pittieri (limón de montaña). A treelet of wet and lower montane forest. Similar to C. indica, but scales are light green or yellowish on leaf under-

side. The fruit is a large round berry. Distribution: Sparse in wet forests, mostly along the Caribbean coast. Recognition: Without flowers, Capparis with evenly spaced leaves are easy to confuse, and in wet forests, there are many species. You have to remember the scaly leaf undersides. Thirteen more species of Capparis are found in Panama, and 10 of those are found in Costa Rica. Three of those are from the dry zones, and at least three more in wet forests, but others are seen too seldom to characterize.

Capparaceae

Capparis indica

Capparis pittieri

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Caricaceae

Crateva tapia

Crateva tapia

Crateva tapia (guaco, palo de guaco, perguetano, mongo). A small tree of dry zones. The trunk is short, forked, and the crown is dense. Bark has white scales. Leaves are alternate, compound, with three leaflets. Distribution: In the driest zones, where it grows in barren, open country. Also on limestone near the Madden Dam, a few coastal sites, and in

swampy areas. Recognition: Easy to learn because there are few trifoliate species, and no others in the dry zones. See species of Allophylus (Sapindaceae), but they have toothed leaves, and Erythrina (Fabaceae—Papilionoideae), but they should not be confused. Capparaceae has two other woody genera, Morisonia and Steriphoma, both with two species of treelets (though Steriphoma is very small). Their leaves are like those of Capparis; Morisonia has small white flowers, but Steriphoma has spectacular large, orange flowers with extended stamens. One species of Morisonia occurs along beaches and mangroves near Panama City and at a few sites in Costa Rica. The others are seldom collected.

Caricaceae A minor family of rainforest trees, with 33 species in the Americas and 2 species in Africa. It includes one of the world’s major fruit crops, the papaya, which grows all over Panama’s lowlands in yards and gardens. There are eight other species in Panama’s tree checklist. Most species in the family have either lobed or palmately compound leaves and milky latex; however, the latex is not abundant in older leaves. All species have extremely soft wood. Carica and Vasconcellea are all small treelets, whereas Jacaratia are tall trees.

Carica papaya (papaya). Not illustrated. The yard fruit. Scars where old leaves have fallen cover the

lower trunk. The leaves are large and deeply lobed, and occur only at the very top. The well-known fruits grow out of the trunk just below the leaves. Male trees produce small flowers on long stalks, but no fruit. Distribution: In yards, gardens, and farms everywhere in the world’s tropics. Recognition: The large leaves are lobed in an intricate way, unlike any other tree but vaguely resembling the epiphytic Phyllodendron (family Araceae, which are not in the book because they are not woody). Papaya is cultivated in origin and might be a hybrid between two naturally occurring species.

Jacaratia spinosa

Jacaratia spinosa

Jacaratia spinosa (papayito). A tall, spiny tree.

The bark spines are sparse and may be missing in older trees. Roots are cylindrical and swollen beyond a constriction where the root meets the trunk. Small branches have many spines. Leaves are palmately compound, with five to nine narrow, pointed leaflets. Breaking a leaf or small branch produces a flow of milky latex. Flowers are white, born on branches; fruits resemble papaya. Distribution: Known in lower montane and Caribbean wet forests from c. Panama to Costa Rica, plus the Osa Peninsula. A single tree is known on Barro Colorado I. Where we know

it, it is very rare, with a few big trees and no juveniles. Recognition: Easily mistaken for Ceiba pentandra or Pachira species (Malvaceae—Bombacoideae), both of which have palmate leaves and trunk spines. But Jacaratia has latex and none of the bombacoids do. Two other species of Jacaratia are known, one in mountains of Chiriqui and widely in Costa Rica in both mountains and wet lowlands. The other is rarely seen. They also have palmately compound leaves. Another genus, Vasconcellea, is closely related to the papaya and was formerly part of Carica, but now only the garden crop gets the latter name. V. cauliflora is nearly identical in appearance to the papaya, but has flowers and fruits growing out of the stem near the ground instead of near the leaves; it grows along the Pacific coast at a few spots through Panama and Costa Rica, and is present but sparse near the Canal. The other two species are only known at a couple places in Panama; one has simple leaves and a baseball-shaped fruit, quite different from papaya.

Caricaceae

Crateva tapia

Jacaratia spinosa

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Celastraceae

Celastraceae Crossopetalum parviflorum

(Including Hippocrateaceae)

Crossopetalum parviflorum

A medium-sized family of tropical and warm temperate zones, including trees, shrubs, and lianas. The burning bush (Euonymus) of North America is in this family. The family Hippocrateaceae is now merged within, and this larger group has 1100 species in the world and 320 in the American tropics. There are 25 tree and shrub species in Panama, and we cover only 1. This is a difficult family to characterize. All have simple leaves, but about half the species in the family have opposite leaves and half have alternate leaves. Twigs typically zigzag between alternate leaves. Flowers are small and always appear in small clusters along branches at or below the leaves. There is no latex, no odor, no conspicuous hairs or odd colors, and flowers are uninspiring; nothing is distinguishing. In the lowlands of Panama, there are few tree species and they are not common.

Crossopetalum parviflorum (limoncillo). A

treelet of wet forest. The small trunk is short and branched near the base. Leaves are simple and op-

posite, sometimes toothed but not always. The small branches are green toward the tip, and somewhat squared in cross section. Flowers are tiny, red, held in clusters from branches at leaf bases. -Distribution: A Caribbean coast species, quite common near the Canal, also at lower montane sites. Always in the forest understory. Recognition: Simple, opposite leaves with teeth are unusual, especially in the lowlands, and weak teeth that are sometimes lacking make for a good character. Check Cassipourea elliptica (Rhizophoraceae), another weakly toothed lowland species with very similar leaves, and also Mollinedia (Monimiaceae). The other Crossopetalum species is montane and mostly known in Costa Rica. Another major lowland genus is Maytenus, with five species in Panama. M. schippii, with shiny, alternate leaves, is widespread in wet and lower montane forests, and frequently seen around the Canal, though it never is common. The remaining genera in the family, with the number of species in each, are Elaeocarpum (one), Cheiloclinium (one), Perrottetia (four), Quetzalia (one), Salacia (seven), Tontelea (one), Wimmeria (one), and Zinowiewia (two). Their variability makes it difficult to provide identifying hints: Cheiloclinium, Quetzalia, Salacia, Tontelea, and Zinowiewia have opposite leaves, whereas Perrottetia and Wimmeria have alternate leaves. Wimmeria and some Perrottetia have toothed leaves, but the rest do not. Moreover, several of those genera have liana species as well as trees.

Chloranthaceae Hedyosmum bonplandianum

Hedyosmum bonplandianum

nent teeth along the margins, and a spicy odor when crushed. Most species have some stilt-roots growing from the trunk near the ground. The opposite leaves and stem sheath suggest Rubiaceae, but Rubiaceae have neither teeth nor a spicy smell.

Hedyosmum bonplandianum (sauquillo, limoncillo). A small tree of montane forest. Low A small tropical family, mostly in montane forest and mostly trees or shrubs, but there are a couple liana species, and a few grow in lowland and even dry forests. There are 80 species in the world and 44 in tropical America. Panama has seven species of trees and shrubs, of which we cover three. The only genus in the New World is Hedyosmum, and it is easy to recognize. At the base of each pair of petioles, there is a sheath wrapping around the stem. Leaves are simple and opposite, have promi-

branches can grow roots that penetrate the soil separately from the main trunk; new branches and roots can divide multiple times, producing a jumbled network at the base of the tree. New root tips exude a sticky mucilage. Leaves are opposite, with small teeth, and between the leaf bases there is a scarlike ridge with either two or four small, pointed stipules on it. Flowers are green; fruits are white berries wrapped in bracts. Distribution: Throughout montane forests of both countries and usually common. Easy to find at Cerro Campana. Recognition: Other Hedyosmum are similar, but

Chloranthaceae

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Crossopetalum parviflorum

Hedyosmum bonplandianum

this species has fewer secondary veins and finer teeth. Toothed, opposite leaves are found in some Siparuna (Siparunaceae), Crossopetalum (Celastraceae), and Cassipourea (Rhizophoraceae), but

those never have sheaths at the leaf bases. All Rubiaceae have opposite leaves and a stipule between the base of the petioles, but they never have toothed leaves.

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Chloranthaceae

Hedyosmum costaricense

Hedyosmum costaricense

Hedyosmum costaricense (sauquillo). A treelet

of the mountains. The trunk base has small stilt-roots. Leaves are opposite and have broad, well-spaced teeth. Secondary veins are numerous, parallel, and neatly arching forward. Flowers are green or white and fruits are white. Distribution: In montane forests of w. Panama through Costa Rica. Recognition: Toothed, opposite leaves with a sheath base distinguish the genus; the neatly arched secondary veins and broad teeth are distinctive of this species.

Hedyosmum mexicanum

Hedyosmum mexicanum

Hedyosmum mexicanum (sauquillo). A small

tree of the mountains. Leaves are opposite, with closely spaced and obvious teeth. The sheath at the leaf base has many tiny filaments resembling a multiforked stipule. Flowers are green and fruits yellow to white. Distribution: Only in far w. Panama and c. Costa Rica at elevations above 2000 m. Recognition: Other Hedyosmum lack the filamentous sheath. The other four Hedyosmum species are very similar in leaf form; three are scarce and only known from a few records in Panama. H. scaberrimum is the one species in the genus seen in lowland as well as montane forest, and it is found widely.

Chloranthaceae

Hedyosmum costaricense

Hedyosmum mexicanum

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Chrysobalanaceae

Chrysobalanaceae Hirtella americana

Hirtella americana

A medium-sized tropical family of trees and shrubs, with 500 species in the world and 400 in the American tropics. It is a widespread family of rainforest, with many common species, and thus one of the major components of most forests. There are 26 species in Panama, and we cover 8. This is not an easy family to characterize or to recognize. Leaves are always simple, alternate, without teeth, and tend to be spaced regularly along branches, in a flat plane, but the latter tendency can be weak. There is no odor and no latex, and branching is irregular and indistinctive, easily confused with Lauraceae or Euphorbiaceae. In the biggest genus, Licania, many species have conspicuously colored leaf undersides due to small, dense hairs, but this is lacking in some Licania and the other genera. Hirtella can usually be recognized by two

tiny stipules at the base of each leaf; however, these are usually lost on older twigs. Licania often has tiny glands or black spots on the petiole where it meets the leaf, but these are inconspicuous and some species lack them. Flowers are conspicuous in Hirtella, but tiny in Licania. Even experienced botanists have trouble with the family, both in recognizing a tree as Chrysobalanaceae and then determining the species once the family is known.

Hirtella americana (camaroncillo, garrapato). A medium-sized tree of Pacific regions. Twigs are densely covered in red hairs. Leaves are simple, alternate, regularly spaced along branches, in a flat plane, shiny above but densely pubescent with red hairs below. There is a pair of narrow stipules at the base of each leaf that can usually be found on newer twigs. Flowers are pink to white, large, on long spikes, and have extended stamens. Fruits are cherry-sized. Distribution: Mostly on the Pacific slope from Panama City sporadically through Costa Rica, but sometimes at wetter sites. Near the Canal, common in forests around Panama City and also near the Caribbean. Recognition: Should be told as Hirtella by the paired stipules. This is the hairiest species, though H. guatemalensis of wet forests (not illustrated) is quite similar.

Hirtella triandra

Hirtella triandra

Hirtella triandra (camaroncillo, camaron, garra-

pato, conejo, chicarrón). A medium-sized old-growth tree. Leaves are simple, alternate, smallish and short, regularly spaced along branches, in a flat plane,

shiny green above. New leaves have black, circular spots. Stipules are thin, pointed, green, and paired on opposite sides of the branch at the base of every leaf. Petioles are short, with light fuzz. Flowers are white, with long pink stamens emerging well above the petals. Distribution: Widespread in the shaded interior of moist, wet, and lower montane forest. Common at Barro Colorado I., where it is one of the dominant old-growth canopy trees. Recognition: If you think of looking for the paired stipules, you will spot the genus. As Hirtella, this species has smooth leaves, with hairs only on the veins underneath; see H. tubiflora.

Chrysobalanaceae

Hirtella americana

Hirtella triandra

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Chrysobalanaceae Hirtella tubiflora

Hirtella tubiflora

Hirtella tubiflora (camaroncillo, camaron). A

small tree of montane forests. It is like H. triandra in leaf form, but leaves are slightly asymmetric at the

base. Distribution: Mostly in lower montane forest but also lowland wet sites, rather sparsely through Panama and Costa Rica; common at Cerro Campana and Cerro Jefe near Panama City. Recognition: See H. triandra. The asymmetric leaf bases are subtle, but work well for identification if you study a lot of leaves. Panama has three more Hirtella species. All have paired stipules and fuzzy leaves and twigs. H. racemosa is a small forest shrub with conspicuous purple flowers, widespread in most forest types, including dry forest.

Licania affinis

Licania affinis

Licania affinis (cenizo, pellejo de sapo). A large

tree of wet forests. Tall, with a cylindrical trunk lacking buttresses; the bark is brown and warty, like a toad (hence sapo in the common name). Leaves are green above but whitish or gray underneath, due to tiny, dense hairs; there are few secondary veins. Leaves also have a tiny pair of glands barely visible on the margin, just above the petiole. Stipules are

nearly always absent, because they fall soon after a leaf emerges. Flowers are tiny, greenish, on terminal stalks. Distribution: Lowland wet forests of the Caribbean slope in c. and w. Panama and n. Costa Rica. Quite common at Santa Rita. Inside mature forest. Recognition: Not like Hirtella. The tiny leaf glands are useful, and combined with leaf coloration and bark lenticels distinguish this species. Leaves resemble those of Lauraceae in form, but Lauraceae never have light-colored undersides. Quite a number of other species have glands at the leaf base (Alchornea and Sapium in the Euphorbiaceae, Hasseltia in the Salicaceae, and Maranthes in this family). Several other Licania have leaves with the whitish or grayish tint underneath, and quite a few other families do as well (a number of Sapotaceae and Myristicaceae, for example).

Chrysobalanaceae

Hirtella tubiflora

Licania affinis

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Chrysobalanaceae Licania arborea

Licania arborea

Licania arborea (raspa, rasca, rascador). A medium-sized tree of dry forests and pasture. Leaves

are wide, nearly round, whitish or light gray below, with many secondary veins. Stipules are usually lost, but there is sometimes a pair of tiny glands near the leaf base. Distribution: Quite common in the driest areas of the Pacific slope, often in farmland and savannah. Recognition: Similar to L. affinis in leaf coloration, but leaves are much rounder, and bark lacks lenticels. Glands are not reliable. The habitat is distinctive, because L. affinis and the other Licania with light leaf undersides all grow inside mature forest in wet or montane zones.

Licania hypoleuca

Licania hypoleuca

Licania hypoleuca (corocillo, garrapato). A forest tree of wet and montane sites. Trunk cylindrical and straight. Leaves are short and wide, triangular, dark

green and smooth above, with striking white undersides (due to tiny hairs). Flowers are tiny and yellow or green. Distribution: Widespread in lowland wet and lower montane zones, forest interior only. Found at Barro Colorado I. and Soberania but not common. Recognition: The leaves are shorter and the undersides more conspicuously whiter than any other Licania. See also Otoba (Myristicaceae) with white leaf undersides, but Otoba has larger leaves and distinctly verticillate branching. From above, leaves of L. hypoleuca look just like those of Oxandra longipetala (Annoncaceae), but Oxandra lacks any white coloration.

Chrysobalanaceae

Licania arborea

Licania hypoleuca

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Chrysobalanaceae Licania platypus

Licania platypus

Licania platypus (sapote, sangre). A tall forest tree. Large trees have small buttresses at the base. The bark is brown, with lenticels. The leaves are simple, alternate, long and narrow, with straight sides, green underneath, lacking the whitish or grayish tint of other Licania. New leaves are red. Where the leaf margin meets the petiole, there is usually a pair of small, slightly elevated glands. The small flowers are like those of other Licania. Distribution: Widely but

sparsely distributed in Costa Rican lowlands; in Panama only near the Canal, where common at Barro Colorado I. and nearby Soberania. Always in mature forest interior. Recognition: Not an easy species to learn, because its leaves are plain and lack any easy character. Resembles Lauraceae in overall leaf arrangement, and leaves might be confused with leaves of some Pouteria species (Sapotaceae), but Pouteria has white latex. If you remember to check for tiny glands, they are useful. Panama has nine more Licania species in its tree flora, but many are restricted to one or a few sites. Seven reach Costa Rica. All are in wet or lower montane forest. Several have light whitish or even reddish leaf undersides, and so resemble L. affinis. It is generally a difficult genus, and any Caribbean forest may have two or more species.

Maranthes panamensis

Maranthes panamensis

Maranthes panamensis (corocito, corozo, palo de gusano). A tall forest tree. Leaves are simple, alternate, fairly regularly spaced on branches, and lack hairs and are thus shiny green above and below. At the base, where leaf meets petiole, there is a pair of small, barely stalked, glands. Stipules are paired, but readily fall and cannot typically be found. Flowers

are white, with many stamens; larger than Licania but smaller than Hirtella. Distribution: Widespread in lowland wet forests of Costa Rica. In Panama only near the Canal, but frequently seen at wet and some lower montane sites there. Restricted to mature forest. Recognition: Not an easy species to recognize. If you notice the glands at the leaf base, they are a good trait. They look like glands in Croton or Sapium (Euphorbiaceae), but those genera are otherwise unlike Maranthes. Chrysobalanaceae contains three more genera in Panama, Couepia with four species and Neocarya and Parinari with one species each. All are very rare. Chrysobalanas icaco is an alien species, sometimes cultivated, typically seen close to the ocean.

Chrysobalanaceae

Licania platypus

Maranthes panamensis

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Clethraceae

Clethraceae Clethra lanata

Clethra lanata

A small family of tropical mountains and warm temperate areas, with just 38 species in one genus. Panama has three species, of which we cover one. With just the one genus, we do not elaborate family characteristics.

Clethra lanata (nance macho, nancito, nancillo,

mameicillo). A medium-sized forest tree of montane and wet sites. Branchlets have fine, reddish hairs. Leaves are simple, alternate, tightly bunched at the end of branches and not lying in a plane; dark green above, but light colored below with a dense coat of soft hairs and yellow veins. Toward the apex, leaves

can have tiny teeth, but these are often inconspicuous or absent. At the very base, the leaf is sometimes curled. Flowers are conspicuous, white, on long straight stalks above the leaves. Fruits are small capsules that split open to reveals seeds. Distribution: Widespread but rather sparse in lower mountains from c. Panama through Costa Rica, also wet lowlands around the Canal and even (surprisingly) in some lowland sites in dry Pacific areas. Common at Cerro Campana and along Pipeline Rd. Often at edges or along roads. Recognition: The bunching of leaves and dense, white hairs underneath should identify this species. The teeth would help but are not reliable. Other species with whitish undersides do not have bunched leaves. The common name indicates the resemblance of leaves to Byrsonima crassifolia (Malpighiaceae), which also has bunched leaves with dense, light, velvety hair underneath, but Byrsonima has opposite leaves and lacks yellow veins. Two other Clethra species are known, but are very rare. Both are montane.

Clusiaceae

Calophyllum longifolium

Calophyllum longifolium

A major tropical family of trees and shrubs, including many epiphytic (trees growing on other trees), and a few herbs. There are 1100 species worldwide and 600 in the tropical Americas. This excludes the Hypericaceae, which has often been merged into the Clusiaceae. Panama has 43 species of trees and shrubs, of which we cover 11. This is a satisfying family to learn because its characteristics are consistent and defining. All species have opposite leaves and latex that is usually yellow or orange, though sometimes white. Leaves tend to be thick and shiny, and secondary veins are parallel and closely spaced. This combination of characters means the family should nearly always be recognized. The only other group with opposite leaves and latex are the Apocynaceae, and these always have dripping white latex. In Clusiaceae, the latex emerges slowly from a broken leaf, producing small yellow or orange droplets.

Calophyllum longifolium (maría, santa maría, calaba). A large and conspicuous forest tree. To 45 m tall, with a straight, cylindrical trunk, only slightly swollen at the base. Bark on adult trees is gray, with clear, vertical fissures. Leaves are simple, opposite, long, with straight sides, and have highly distinctive venation; secondary veins are extremely densely spaced, straight and parallel, and no smaller veins are visible. Saplings have no branches, so very large leaves emerge from the stem. Breaking a leaf reveals small droplets of yellow latex, appearing slowly and not dripping. Flowers are fairly large, white, with yellow stamens. Fruits are plumsized, round. Distribution: Widespread in moist, wet, and lower montane forests, and common everywhere around the Canal. Mostly in mature forest. Recognition: Leaves of Calophyllum are unmistakable. Only the genera Manilkara and Micropholis (Sapotaceae) might be confused, but they have alternate leaves, not opposite. Large trees of Calophyllum are easy to recognize from the fissured bark, and fallen leaves can always be found on the ground below big trees. Check the other species of Calophyllum in the area, all of which have the same leaves (as do species in Asia and Australia, which are also easy to recognize). María is a well-known species; the wood is used locally for timber, but it remains very common.

Clusiaceae

Clethra lanata

Calophyllum longifolium

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Clusiaceae

Calophyllum nubicola

Calophyllum nubicola

Calophyllum nubicola (maría enano, santa maría, calaba). A small tree of cloud forest. Trunk often branches near the ground; bark has yellow lines. Leaves are short and oval in shape, with finely spaced secondary veins like other Calophyllum.

Distribution: Known only in lower montane forest of c. Panama. Fairly common at Cerro Jefe near Panama City. Recognition: Easy to spot as Calophyllum. It has shorter, rounder leaves than the much more widespread C. longifolium. The two might be seen together in lower montane forests. Two more Calophyllum species are native to Panama. C. brasiliense, a tall tree with yellowish patches in the bark, is widespread and better known in Costa Rica than C. longifolium, but much less common in the Canal Area. The other native species is rarely seen. An Asian species, C. inophyllum, is an ornamental in Panama City and the Canal Area, and any Calophyllum in urban areas is most likely C. inophyllum.

Clusia pratensis

Clusia pratensis

Clusia pratensis (copé). A treelet of cloud forests. It can grow on the ground, but also on other trees. On the ground, the short trunk branches often and sometimes has stilt-roots. Leaves are roundish, very thick, secondary veins are barely visible. Broken leaves or bark produce cream- or yellow-colored latex that drips slowly. Flowers are large and pink; fruits are a woody capsule that splits open into a starshape with seeds embedded under the arms. Distri-

bution: In lower montane forest of c. and w. Panama. Common at Cerro Campana. Recognition: Leaves of Clusia are curiously thick and very distinctive. There are more species that grow as epiphytes, and leaves can often be found on the ground where the Clusia itself grows far above and out of sight. One species in the Rubiaceae, Cosmibuena macrocarpa, has similar leaves, also opposite, but without latex. Cosmibuena also grows in montane forest, and if you don’t look for latex, you will confuse it with Clusia. Ten species of Clusia are found in Panama, and all have similar leaves, fruits, and flowers. C. stenophylla is the most widespread and commonly seen species in both Costa Rica and Panama. Clusia species in general are most common in montane forest, whether growing on the ground or as epiphytes. In lowland forest, they are exclusively epiphytes and seen mostly as leaves on the ground.

Clusiaceae

Calophyllum nubicola

Clusia pratensis

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Clusiaceae

Garcinia intermedia

Garcinia intermedia

Garcinia intermedia (madroño, chaparrón, sastra, sastro). A medium-sized tree of forests throughout Panama. Not quite (or just barely) reaching the canopy, with a straight trunk. Branching is notable, with groups of three or four arising together, and these groups regularly spaced up the trunk. Leaves are thick, glossy green, dark green above and lighter below, opposite, oval, long, with straight sides, and parallel secondary veins that are faint but visible. Breaking a leaf or twig reveals a bit of whitish or

cream-colored latex. At the end of a branch, between the final leaf pair, there is a crack that spreads when the two leaves are pulled apart; this is a good character of the genus Garcinia everywhere in the world. Flowers are fairly large, whitish yellow, born at leaf bases. Fruits are smooth. Distribution: Widespread in wet to moist forest and also lower montane and even fairly dry sites, only inside mature forest. Common in old growth at Barro Colorado I. Recognition: The genus Garcinia is easy to learn from the opposite leaves and faint but clear secondary veins, even without checking for latex. But this and the next species, G. madruno, are very difficult to tell apart; the only clear difference is latex (cream-colored in G. intermedia, yellow in G. madruno), but even that is slight. Symphonia is also similar, but has narrow, pointed leaves, lacks the gap between the end pair, and has very distinctive roots. See also leaves of Aspidosperma spruceanum (Apocynaceae), which look just like leaves of Garcinia but are alternate, not opposite.

Garcinia madruno

Garcinia madruno

Garcinia madruno (madroño, chaparrón, sastra, sastro). A medium-sized tree of wet or moist forests. See the very similar G. intermedia, but this species

has yellow latex and warty fruits. Distribution: More a Caribbean wet-forest species than G. intermedia, and does not occur in dry regions. Like that species, it is common in old growth at Barro Colorado I. and restricted to mature forest. Recognition: See G. intermedia. Three other species of Garcinia in Panama resemble the two presented, but they are seldom seen. The mangosteen (G. mangostana), native to Southeast Asia, is frequently planted and fruits can be found at many markets; there is a big stand at Summit Garden near the Canal.

Clusiaceae

Garcinia intermedia

Garcinia madruno

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Clusiaceae

Marila laxiflora

Marila laxiflora

Marila laxiflora. A small tree of wet and moist forest. Trunk is short and ordinarily has low branches. Leaves are simple, opposite, thick and shiny, and with distinct venation: the tertiary veins (the smallest visible) emerge from the secondary veins, but curve toward the center to meet the primary vein; only those near the center do this, and it is only visible on

the leaf underside. Each secondary vein loops forward near the leaf margin to meet the next secondary. Leaves have tiny clear spots, visible only with magnification. Petiole is twisted and has a channel above. Broken leaves, branchlets, or bark ooze milky latex that slowly forms drops. Flowers are large, whitish to green. Fruit is a long, thin capsule, opening to produce seeds with tiny cottony hairs at both ends. Distribution: Wet and lower montane forests throughout, reaching moist sites, including Barro Colorado I. Only inside mature forest. Recognition: The shiny, opposite leaves, with latex, match the typical Clusiaceae pattern. The venation is distinctive of the genus. See details of venation and the petiole of the very similar M. pluricostata. The family Myrtaceae has opposite leaves and clear dots in the leaves, but lacks latex.

Marila pluricostata

Marila pluricostata

Marila pluricostata. A small tree of wet and montane forest. See the description of the tertiary veins of M. laxiflora, which this species shares. Leaves are long and straight-sided, have numerous

nonarching secondary veins that are reddish or yellowish (on the underside). The petiole is deeply channeled and twisted. Flowers are pink and fruits are like those of M. laxiflora. Distribution: More restricted to wet and lower montane sites than M. laxiflora, not reaching moist forest and not at Barro Colorado I. Only in mature forest. Recognition: The longer leaves and petiole best distinguish this species from M. laxiflora. Two other species of Marila are rare and known only in c. Panama. Both have the same distinctive venation.

Clusiaceae

Marila laxiflora

Marila pluricostata

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Clusiaceae

Symphonia globulifera

Symphonia globulifera

Symphonia globulifera (cerillo, cero, barillo). A tall tree of swamp forest. Trunk is straight but has numerous long stilt-roots at the base emerging well above the ground. Leaves are simple, opposite, narrow and pointed, with faint secondary veins. Any

broken part produces bright yellow latex. Flowers are small, red globes that can be found on the ground under big trees. Distribution: Widespread in Caribbean lowlands, lower montane sites, and the Osa Peninsula, just reaching moist forest at Barro Colorado I. In Bocas del Toro, grows in swamps near the mangroves, but occurs in upland forests widely too. Recognition: The shiny, opposite leaves are conspicuously Clusiaceae. Compared to the similar Garcinia, this species has small, narrow, pointed leaves; recall also that Garcinia has a narrow slot between the end pair of leaves which Symphonia lacks. Even without seeing leaves, any really large tree with numerous stilt-roots should at least bring to mind Symphonia.

Tovomita longifolia

Tovomita longifolia

Tovomita longifolia. A small tree of wet and montane forest. Trunk has low branches, and at its base there are many long stilt-roots. Leaves are simple, opposite, shiny, thick, smooth, and bunched toward the end of branches. Breaking off a leaf produces small drops of yellow latex. Flowers are

medium-sized, with rounded white to yellowish petals. Fruits are capsules. Distribution: Wet and moist to lower montane forest throughout; found at Soberania and nearby areas around the Canal, but not especially common. Inside mature forest only. Recognition: It is unlike other Clusiaceae in having leaves bunched toward branch tips rather than spaced evenly. The thick, smooth leaves are also distinct. Prominent secondary veins are most like those of Marila, but the latter has distinctive venation. A species of Rubiaceae, Tocoyena pittieri (not illustrated; see Posoqueria latifolia), has similar opposite, shiny leaves, but all Rubiaceae have a stipule between the petioles and lack latex. See the other Tovomita, all with thick, shiny leaves.

Clusiaceae

Symphonia globulifera

Tovomita longifolia

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Clusiaceae

Tovomita stylosa

Tovomita stylosa

Tovomita stylosa. A treelet of forest interior. Similar to the previous species in all respects, with thick shiny leaves and yellow latex. But leaves are much

smaller and pointed at the tip. Flowers have long, narrow, greenish petals and greatly extended stamens. Fruits are small capsules. Distribution: Widespread in wet, moist, and lower montane forest through Panama just to s. Costa Rica. Inside mature forest only. Recognition: Small leaves distinguish this easily from T. longifolia, but also mean it is confused with other opposite-leaved species with small leaves. Tabernaemontana arborea (Apocynaceae) has similar leaves, but drips white latex. Several Psychotria (Rubiaceae) have small, opposite leaves, but never have latex, and the leaves are not as thick and smooth.

Tovomita weddelliana

Tovomita weddelliana

Tovomita weddelliana. A small tree of wet and montane forest. Trunk and stilts are just like those of the previous two species. Leaves are thick, smooth, shiny, very long and narrow, widest close to the tip, and bunched densely toward the end of branches; secondary veins are barely visible. Distribution: Montane and lowland wet forest of the Caribbean slope and the Osa Peninsula, more restricted than the previous two species and not reaching moist for-

est. Recognition: The thick, long leaves are easily recognized. A few odd little trees have long, thick leaves, including Euphorbia elata (Euphorbiaceae) and Potalia amara (Loganiaceae), but they are rare (none are covered in this book) and much different from T. weddelliana. Clusiaceae has three more genera, all with opposite leaves and latex. Chrysoclamys has 10 species; C. eclipes is widespread from c. to e. Panama and common in forests near the Canal; C. glauca is most widespread in Costa Rica. Leaves are rather thick and smooth, held at various angles from the branches. One Dystovomita species is restricted to high elevations and has thick, rounded leaves with prominent veins. Mammea has a native species with smooth, green, narrow leaves, known only in lowlands near the Canal, plus M. americana, the mamey, an American species cultivated for fruits.

Clusiaceae

Tovomita stylosa

Tovomita weddelliana

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Cochlospermaceae

Cochlospermaceae Cochlospermum vitifolium

Cochlospermum vitifolium

A tiny, tropical family of small trees, with 12 species worldwide. Asian species are used as ornamentals in other parts of the world. The group is sometimes combined with the Bixaceae. There are two species in Panama. We omit family traits for such a small group.

Cochlospermum vitifolium (poro-poro). A

small tree of grasslands. Leaves are simple, alternate,

toothed, with pointed lobes, reminiscent of North American maples (Acer). Flowers are large, yellow, showy, conspicuous from early December throughout the dry season. All leaves are dropped while flowering. Fruits are green capsules, and seeds within are tiny, with cottony filaments. Distribution: Grows nearly anywhere there is not forest: gardens, abandoned farms, hedgerows, and wasteland. One of the few trees growing among tall grasses that burn regularly, the poro-poro is remarkably resistant to fire. Recognition: The palmately lobed leaves are easy to recognize in its grassland habitat, even before the big yellow flowers appear. Another species of Cochlospermum, C. orinocense, is known rarely in e. Panama. Its flowers are similar, but the leaves are palmately compound, with five to seven leaflets (like Pseudobombax septenatum [Malvaceae—Bombacoideae]).

Combretaceae Buchenavia tetraphylla

Buchenavia tetraphylla

have consistent traits that make them fairly easy to learn. The leaves are short, widest near the tip, and bunched toward the end of branches, indeed, sometimes they are so bunched it is difficult to tell whether they are alternate or opposite. Flowers are small, white, and held on spikes.

Buchenavia tetraphylla (amarillo, amarillo de A modest family of trees, lianas, and occasionally small shrubs, nearly all tropical, with about 500 species in the world and 85 species in tropical America. There are eight species of trees in Panama, of which we cover six; many more are lianas. The six species we cover here are tall, straighttrunked trees. Two are mangroves and not like the rest, but can be identified by their seashore habitat. The remaining four species, the non-mangroves,

pepita). A tall and rare tree. Trunk is straight, can be large, with medium-sized buttresses at the base. Leaves are small, densely bunched at the end of branchlets, and distinctly widest close to the tip. Flowers are white, on spikes. Fruit is fleshy, one-seeded. Distribution: Known only at a few sites along the Pacific slope plus near the Canal. Recognition: See the similar and far more common Terminalia amazonia. Buchenavia has smaller, narrower leaves, and in case you find fruits, those of Terminalia are quite different.

Combretaceae

Cochlospermum vitifolium

Buchenavia tetraphylla

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Combretaceae Bucida buceras

Bucida buceras

Bucida buceras (mareon, amarillo). A large tree of the Caribbean coast. Trunk is straight and can be large, with swollen roots at the base. The bark is brown, ridged, peeling in small pieces. Branchlets

near leaves sometimes have small but sharp spines. Leaves are shiny green, short, rounded, often with a V-shaped indentation at the tip, broader closer to the tip than to the base. Flowers are white, on spikes, and fruits are small nuts carrying remnants of the flower. Distribution: Along the Caribbean coast in Panama only, usually in swamps but not in salt water; fairly easy to find at Fort San Lorenzo at the mouth of the Chagres River. Recognition: The rounded, bunched leaves resemble those of the widespread Terminalia amazonia, as does the bark, but the latter has large plank buttresses and never has spines, and seeds of Bucida are not winged.

Conocarpus erectus

Conocarpus erectus

Conocarpus erectus (mangle botón, mangle botoncillo, button mangrove). A small tree of beaches. The trunk is short and branches are low. Leaves are simple, alternate, with two small glands at the base.

There are small pits along the primary leaf vein. Individual flowers are tiny and green, held in a dense ball. Fruits are likewise segmented balls with short warts; they float and are dispersed in water. Distribution: Grows on beaches or the land side of mangrove forests, on both coasts throughout. Recognition: If you spot this as a mangrove, it is easy to recognize because it is the only one with alternate leaves. But because it does not grow inside mangrove swamps, it may not immediately be recognized as a mangrove, and there are other beachside trees that are not mangroves. The narrow, pointed leaves with clear pits along the main vein are distinctive.

Combretaceae

Bucida buceras

Conocarpus erectus

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Combretaceae

Laguncularia racemosa

Laguncularia racemosa

Laguncularia racemosa (mangle blanco, white mangrove). A true mangrove. The trunk is straight

and cylindrical, and the bark gray with vertical fissures that deepen in large trees. Roots extend away from the base of the trunk near the surface, and pneumatophores emerge from the roots above the mud. There are not stilts at the trunk base. Leaves

are opposite, rounded at both ends, smooth, with secondary veins barely visible. On the petiole just below the leaf are two small, round, flat glands. Flowers are white, in spikes emerging between leaves. Fruits are ribbed capsules that float. Distribution: In saltwater swamps on both Caribbean and Pacific coasts. Recognition: You only need to separate this from the other mangroves, because no other trees grow in saltwater swamps. Most of the mangroves have opposite leaves, but of those, only Laguncularia has small glands on the petiole. Conocarpus also has glands but its leaves are alternate and otherwise much different. Avicennia (Acanthaceae) is most like Laguncularia, but lacks glands and has narrower leaves that are much lighter below than above. The other mangrove, Rhizophora (Rhizophoraceae), has stilt-roots.

Terminalia amazonia

Terminalia amazonia

Terminalia amazonia (amarillo, roble amarillo,

amarillo carabazuelo). A tall, stately tree of secondary forest. Trunk is straight, with substantial plank buttresses in big trees. Branches and leaves form flattened layers up the crown, pagoda-like; this branching pattern is even called terminalian. Bark is brown, heavily fissured. Leaves are alternate, bunched toward the end of branches, short, widest near the tip and narrowing abrubtly to a little point. The few sec-

ondary veins arch forward toward the tip. Flowers are small, white, on long spikes. Fruit is a tiny seed with two little wings. Distribution: Widespread in moist, wet, and lower montane forests. Abundant in secondary forest throughout the Canal Area. Does not invade roadsides or fields, but establishes when forest is young. In 60- to 80-year-old forests around Panama City, this is often the one big, buttressed tree. Recognition: Its abundance in secondary forests makes it fairly easy to learn. The brown fissures and buttresses on big trees in otherwise small forest stand out, as does the branching form. In juveniles, the short leaves that are broadest near the tip are distinctive; note that other Combretaceae have similar leaves, but none of the other species are anywhere near as common. There are species of Psychotria (Rubiaceae) whose leaves are shaped just like those of T. amazonia, but the Psychotria have opposite leaves and are at most a few meters tall.

Combretaceae

Laguncularia racemosa

Terminalia amazonia

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Cornaceae

Terminalia oblonga

Terminalia oblonga

Terminalia oblonga (guayabo de montaña, guay-

abillo, guayabón). A tall, uncommon forest tree. Trunk is straight, cylindrical, with high branches, and bark is smooth, cream or yellow colored, and has large, gray sheets flaking off. Leaves are alternate, larger than in T. amazonia, wider slightly nearer to the tip, often but not always bunched toward the ends of branches. Flowers are small, white, on long spikes. Fruits have pairs of wings, small but slightly larger

than T. amazonia. Distribution: Occurs throughout, from fairly dry to wet and lower montane forest, but nowhere near as common as T. amazonia. Recognition: The leaves resemble those of other Combretaceae in shape and bunching, but less consistently so and are sometimes confusing. In big trees, the smooth, peeling bark is distinctive. The common name demonstrates the similarity to the guava tree (Psidium guajava [Myrtaceae]), which also has smooth, light bark, but guava is a small tree with opposite leaves and will not be confused. The one other native Terminalia species is known from mangrove swamps and is very rarely seen. In addition, a common ornamental from Asia, T. catappa, is planted throughout cities and spreads on its own along both ocean coasts. Its leaves are clustered, alternate, widest near the tip, but much larger than those of the native Terminalia.

Cornaceae Cornus disciflora

Cornus disciflora

A small family of about 100 species worldwide, mostly in the dogwood genus, Cornus, of the north temperate zone. In tropical America, there are just four species, only in mountains of Central America and the northern Andes. Because there is just one species in Panama, we omit a family description.

Cornus disciflora (mata hombro, lloró). A tall,

montane tree. Trunk is straight, cylindrical, with bark flaking in small pieces. Leaves are simple, opposite, and have a bluish green underside. The few secondary veins arch well forward toward the leaf tip. Flowers are small, greenish to yellow, in small heads. Fruits are single-seeded, fleshy. Distribution: Only in high mountain forests, above 2000 m, in far w. Panama through Costa Rica. Recognition: If you know the North American dogwoods, you will recognize this; the leaves are alike. Viburnum (Adoxaceae) of the same habitat is rather similar. Cornus also might be confused with Rubiaceae, which have simple opposite leaves, but Rubiaceae have stipules or stipule scars across the branch between leaf pairs.

Cornaceae

Terminalia oblonga

Cornus disciflora

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Cyrillaceae

Cyrillaceae Cyrilla racemiflora

Cyrilla racemiflora

A tiny family of tropical and warm temperate areas, with just 14 species in three genera worldwide and 12 species in the Americas. Panama has one species.

Cyrilla racemiflora (mangle cimarrón). A small

cloud-forest tree scarce in Panama. Leaves are simple, alternate, bunched toward the ends of branches, narrow, thick and smooth. Secondary veins are small, inconspicuous, forming a fine network. Old leaves turn orangish, and most trees have some colored leaves. Flowers are white, in long spikes below the leaves. Fruits are small berries. Distribution: Known only in cloud forests in c. Panama, and we recently also found it in swamps near the Caribbean coast in Bocas del Toro. Recognition: The smooth, thick leaves are quite distinctive. They are rather like those of Morella (Myricaceae) of cloud forests.

Dilleniaceae Curatella americana

Curatella americana

A moderate-sized tropical family, almost entirely lianas. There are 335 species in the world and 70 in tropical America. In the New World, only a few species are small shrubs, and just one is a tree, which we cover here.

Curatella americana (chumico, chumico sabanero). A small, bent tree of the region’s driest savannahs. Trunk is typically short, irregular, branched near the ground. Leaves are alternate, with wavy margins, sandpaper-rough to the touch. Flowers are pinkish, in dense terminal clusters. Fruits are capsules that split open to reveal black seeds with fleshy, white arils. Distribution: In dry open country of the Pacific slope, where it can be numerous. Never in forest. Recognition: The cashew, Anacardium occidentale (Anacardiaceae), has similar form and grows in the same habitat, but its leaves are smooth. The cashew is also distinguished by having a few orange leaves, never present in Curatella.

Dilleniaceae

Cyrilla racemiflora

Curatella americana

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154

Ebenaceae

Ebenaceae

Diospyros artanthifolia

Diospyros artanthifolia

The ebony family is a medium-sized group of 500 tree species, including 100 in the Americas. A few species occur in subtropical or warm temperate areas, including the persimmon (Diospyros kaki), but nearly all are tropical. There are seven species known in Panama, all in the genus Diospyros, and we cover one. Ebony is known for its hard, dark wood, but not all species share this character. Apart from the wood, they are not easy to recognize. Leaves are simple, alternate, and arranged regularly in a single plane. Groups of branches emerge from a single point on the trunk, creating regular layers. In both those traits, Ebenaceae resembles Annonaceae. Leaves of Diospyros, however, tend to be hairy, and the one species we cover here has dense hairs.

Diospyros artanthifolia (sapote negro). A me-

dium-sized forest tree. Trunk is straight and cylindrical, and bark is black. The outer branches are long, thin, and covered with dense reddish hairs. Leaves are simple, alternate, regularly arranged in a plane, and have dense reddish hairs, especially on the veins. There are many small black spots on the leaf underside that should be visible without magnification. Flowers are white, in leaf axils. Fruits are large, yellow, and fleshy. Distribution: Only known on the Caribbean half of the Canal Area and a little ways east. Not common, and never outside the forest. Recognition: The leaf arrangement resembles small-leaved Annonaceae, especially Guatteria dumetorum, but Annonaceae lack hairs. Look for the long, thin branches and black leaf spots of Diospyros.

Diospyros has six more species, all poorly known in Panama. Two are better known in Costa Rica (D. digyna and D. hartmanniana). Without the fruit, they are not easy to recognize; some but not all have black spots or glands on leaves, and several but not all have lots of pubescence.

Elaeocarpaceae Sloanea terniflora

Sloanea terniflora

extend well above the calyx. Nearly all Sloanea have capsular fruits covered in long, twisting or curving spines (like Lindackeria [Achariaceae]); however, the one species we cover has very short spines.

Sloanea terniflora (terciopelo, mameicillo, casa-

A medium-sized tropical and Southern Hemisphere family of trees and shrubs, with 500 species worldwide and 100 in tropical America. Sloanea is the only major genus in the New World tropics, and the only one in Panama. Here we cover 1 of the 11 species in Panama’s tree checklist. Sloanea species are all medium or large trees, with simple, alternate leaves; a few have toothed leaves, but most not. A good character for the family is the petiole, which is swollen at both ends; moreover, the leaf is held at an angle relative to the petiole. We call this petiole arrangement a doubleelbow. If these swellings were green, they would resemble those of Fabaceae, but they are brown; some Malvaceae have petioles swollen at both ends, but they have palmate venation. Flowers have no petals but a rather thickened calyx that might look like the petals; there are many stamens that often

co). A large forest tree. Trunk can be huge, with smooth gray bark and many irregular, rounded buttresses at the base. Leaves are simple, alternate, bunched toward the end of the branches, but highly variable in other characters, such as rounded or pointed, toothed or not, sometimes hairy; juvenile leaves especially have hairs and can be toothed. The consistent character is the double-elbow and the angle between leaf and petiole. Flowers are brown or red with numerous stamens forming a dense brush. The fruit capsule is covered in very short spines. Distribution: Most widely found in the driest areas, in c. Panama and nw. Costa Rica, and it can be found regularly along streams in this region. Also at a few wetter sites along the Pacific coast and the Caribbean side of the Canal. Fairly common in secondary forest around Panama City and old growth at Barro Colorado I. Recognition: The leaves bunched toward branch ends somewhat resemble Terminalia and other Combretaceae. Leaf shape is highly variable, though. In big trees, the buttresses are distinctive.

Elaeocarpaceae

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Diospyros artanthifolia

Sloanea terniflora

Sloanea is a diverse genus with many locally distributed species. At least 10 more occur in Panama, 7 of which reach Costa Rica. S. zuliaensis is found in the Canal Area, at Barro Colorado I. and Soberania; it has a spiny, green fruit and bluntly toothed leaves. Nearly all the other species are restricted to

wet and lower montane forest; some are poorly known. Our surveys uncovered two species not previously known in Panama, and possibly several more we have not yet identified. Sloanea can reliably be recognized by the double-elbow and angled leaf.

156

Erythropalaceae

Erythropalaceae Heisteria acuminata

(Formerly Olacaceae)

Heisteria acuminata

that nevertheless proves easy to learn. Flowers are tiny and inconspicuous.

Heisteria acuminata (sombrerito, ajicillo, chorola). A forest treelet. See the description of the more

A small tropical family of primarily trees and shrubs, though some are lianas. It was until recently classified within the Olacaceae. There are somewhere around 100 species worldwide and 40–50 in the American tropics. Here we describe 3 and illustrate 2 of the 11 species known in Panama. Erythropalaceae species have simple, alternate leaves, and often have latex, though not much. Most species in Panama, and all those we cover here, have sharply upturned petioles, a seemingly subtle trait

common Heisteria concinna, which this species resembles in most features, including the Mexican-hat fruit. But H. acuminata is a smaller plant, with smaller leaves, and with petioles less distinctly curved. Distribution: Widespread from moist and wet to lower montane sites, especially on the Caribbean slope. Fairly common in the Canal Area, more so toward the Caribbean. Restricted to forest; not known in open areas. Recognition: Quite similar to H. concinna, but this species is more difficult to recognize because the distinctive features are less pronounced. The petioles curve upward less clearly than in H. concinna, and the branches droop less. As saplings, H. acuminata has much smaller leaves, and the upturned petiole remains a good clue.

Heisteria concinna

Heisteria concinna

Heisteria concinna (sombrerito, ajicillo, chorola,

Mexican hat). A medium-sized forest tree. Trunk is straight and cylindrical, and the bark black with lenticels. Branches are long and hang downward, resembling liana branches. Leaves are simple, alternate, shiny green and smooth above, and arranged regularly along branches. Sapling leaves in the understory are long and straight-sided; canopy leaves are smaller and closer to triangular. The petioles are U-shaped, curving upward conspicuously, and are swollen toward their apex. Broken leaves produce a small amount of white latex. Flowers are tiny, in stalkless clusters at the base of leaves. Fruits are berries

that turn white, on top of a circular reddish bract at the base (a remnant of the flower’s calyx); because of the red base, they are often mistaken for flowers. Together the small white berry and reddish base look like a little hat, hence the common names. Distribution: Widespread in lowland forests throughout, not usually in mountains and known fairly widely along the Pacific coast. Very common in the Canal Area, where it is often seen as a sapling in mature forest on Barro Colorado I. and secondary forests near Panama City. Does not grow in open areas. Recognition: The long, hanging branches are easy to spot with some practice. Shiny leaves with an upturned petiole are also distinctive. See H. acuminata. Five more tree species of Heisteria are found in Panama. All are restricted to the Caribbean slope or montane forests. Most have upturned petioles. H. costaricensis is shrub-sized and has peculiarly long, very narrow, pointed leaves; it is found near the Caribbean along the Canal and at a few widely scattered sites in Costa Rica.

Erythropalaceae

Heisteria acuminata

Heisteria concinna

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Erythroxylaceae Minquartia guianensis

Minquartia guianensis

Minquartia guianensis (cuajado, cuajado negro, criollo, criollo negro). Not illustrated. A tall wet-forest tree. Trunk is extremely fluted, and the gray bark peels off in small pieces. Leaves have straight, parallel secondary veins, and very finely spaced and par-

allel tertiary veins running at right angles to the secondary veins. Petioles are sharply upturned. Broken leaves slowly produce white latex. Flowers are in long spikes; fruits are ovoid and fleshy (not like hats as in Heisteria). Distribution: In Panama, known only in lowland Caribbean forests near the Canal, where it is not especially common. In Costa Rica, found on the Osa Peninsula and in Caribbean lowlands. Recognition: The fluted trunk can be confused with Aspidosperma megalocarpon (Apocynaceae), Macrocnemum roseum (Rubiaceae), or Platypodium elegans (Fabaceae—Papilionoideae). A close look at the leaf venation should help. The upturned petioles resemble those of Heisteria, and also vaguely Sloanea (Elaeocarpaceae).

Erythroxylaceae Erythroxylum citrifolium

Erythroxylum citrifolium

A modest-sized family of tropical treelets and shrubs, with 250 species worldwide and 180 in tropical America. All the American species are in the genus Erythroxylum, which is the coca genus, and several species have psychoactive alkaloids in their leaves. Cocaine is derived from Erythroxylum coca, a South American species. Panama has eight species in the genus, of which we cover three. All are understory treelets, mostly of mature forest, with simple, alternate leaves. The best family character is the presence of brown stipules. These are pointed sheaths, often numerous and persisting on many branches and thus conspicuous. In addition, leaves of Erythroxylum often have a light green

band on the underside, parallel to the midrib. Sometimes this is very clear, but sometimes it is absent or barely visible. Flowers are very small, produced on branches usually at leaf axils; fruits are red berries.

Erythroxylum citrifolium (alcarreto). A small tree or treelet of moist and wet areas. Trunk is straight but short. Leaves are simple, alternate, narrow and pointed, shiny green, a dark shade above but light and yellowish below. The leaf underside lacks any central light band, but small veins form a fine network. The stipules are brown, long and pointed, and persistent. Flowers are tiny, in little groups at leaf clusters. Fruits are elongated red berries held on the branches. Distribution: Widespread all around the Canal Area, but only known at a few other widely scattered sites in Panama and Costa Rica. Seldom common. Can be found pretty easily along forest edge at Santa Rita. Recognition: Leaves resemble those of several Annonaceae, and could easily be confused. Look for the pointed brown stipules, which are nearly always present, and the fine network of tertiary veins.

Erythroxylum macrophyllum

Erythroxylum macrophyllum

Erythroxylum macrophyllum (alcarreto). A small tree or treelet of forest interior. Leaves are simple, alternate, rather bunched toward the end of the primary stem or branches, shiny dark green above.

Below they are an attractive yellowish green with finely reticulate tertiary veins. There is a vaguely brighter section along the midrib. On the stem or branches around the leaf bases are large, papery, brown stipules, numerous and very conspicuous. Flowers and fruits are like those of E. citrifolium, but slightly larger. Distribution: Widespread in montane, wet, and moist forest throughout, but never especially common. Recognition: The dense brown stipules are unmistakable and should clue you in to looking closely at the leaf undersides. Leaves of E. citrifolium are quite similar, but do not have such large and obvious stipules.

Erythroxylaceae

Erythroxylum citrifolium

Erythroxylum macrophyllum

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Euphorbiaceae

Erythroxylum panamense

Erythroxylum panamense

Erythroxylum panamense (alcarreto). A small

tree or treelet of forest interior. Leaves are fairly small, narrow, pointed, shiny green, with a very clear light band on the underside along the midrib. Brown stipules are pointed, persistent, but small. Flowers

and fruits are like those of other Erythroxylum. Distribution: Known from c. Panama east. Not common but found many places near the Canal. Recognition: The light bands on the leaves are pronounced and distinctive. If you don’t check the underside, the leaves look quite a bit like those of Ouratea lucens (Ochnaceae), though the latter has teeth and much different venation. The remaining four Erythroxylum species are very seldom seen. Leaves are quite variable in form, and not all species have conspicuous stipules or the light band on the leaf underside, so are difficult to recognize.

Euphorbiaceae Acalypha diversifolia

Acalypha diversifolia

A large family, one of the major tropical rainforest families throughout the world. There are 7000 species worldwide, found in temperate, tropical, and desert climates; tropical America has 1500. Included within the family are two species of great economic value internationally: Brazilian rubber (Hevea brasiliensis) and manioc (Manihot esculenta). Both are native to Amazonia but are often cultivated in Panama. Euphorbiaceae is also the family of the castor bean, which grows widely as a weed in the tropics and is sometimes cultivated. Among native euphorbs, Panama has 69 species of trees and shrubs and many more vines and herbs. We cover 16 species here. There are two other families recently segregated from the Euphorbiaceae: Phyllanthaceae and Putranjivaceae, which we treat separately. The Euphorbiaceae have always been infamously variable. Even after segregating the Phyllanthaceae and Putranjivaceae, the species are diverse, and the family does not characterize well. They have alternate leaves, sometimes compound or lobed but usually simple. There are small glands at the base of the petiole in quite a few, and many have milky latex, but many species lack those traits. In a few species, the latex is toxic. There is one section of euphorbs, however, that can usually be spotted by the leaves: they have broad, rounded, heart-shaped leaves

bunched toward the end of branches, with the leaf angled from the petiole. Many in this group routinely carry a few yellow or orangish leaves in their crown, a surprisingly useful tidbit. Many euphorbs have small to tiny flowers on dense spikes, but others break this pattern. Most have fruits in dry capsules that split open to reveal seeds. Any tree or shrub with rounded or heart-shaped leaves, and any with small glands at the petiole base, should at least call to mind Euphorbiaceae.

Acalypha diversifolia (palito feo, prende-prende). A scraggly and weedy shrub that stretches our definition of tree. The small stems branch frequently and are leaning or arching. Leaves are toothed, but variable in shape. Flowers are on a small spike at leaf axils; individual flowers are nearly microscopic. Fruits are very small capsules with three sections. Distribution: Found everywhere in Panama and Costa Rica, in forest, woodlots, and roadsides. In wetter areas can be abundant; along Pipeline Rd. in Soberania, for instance, it forms a dense, weedy hedge in some areas. Also grows inside mature forest where there is some light. Recognition: Know this little weed from the arching stems along with toothed leaves. The leaf shape is tricky, since it is variable and can be similar to that of a number of other species. Note the various Salicaceae, for instance, with toothed leaves, and Hybanthus prunifolius (Violaceae) is also similar but has white stipules. Three other Acalypha species are found in Panama. One of them, A. macrostachya, is less common in similar habitat to A. diversifolia. Its leaves are heartshaped and flowers are tiny and red. The other two species are scarcely seen in Panama; A. villosa is better known in Costa Rica.

Euphorbiaceae

Erythroxylum panamense

Acalypha diversifolia

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Euphorbiaceae Adelia triloba

Adelia triloba

Adelia triloba (espino amarillo, bagre). A mediumsized forest tree. The trunk often has low branches and sometimes small stems sprouting near the base; in bigger trees, the trunk is fluted. There can be

spines on small branches. Leaves are simple, alternate, have few secondary veins, and lack teeth. Each leaf grows on a small, woody swelling off the side of the branch, not directly from the branch. Flowers emerge from tiny clusters along the branches, among the leaves. Fruits are capsules with three sections. Distribution: Known widely in moist lowlands and some drier areas, and on limestone outcrops near the Canal. Never very common. Recognition: The small woody base to each leaf is the trick; spines are also useful, because few forest trees have them. Leaf shape resembles that of Acalypha diversifolia and Hybanthus (Violaceae), but both have teeth.

Alchornea costaricensis

Alchornea costaricensis

Alchornea costaricensis (nagua blanca). A medi-

um-sized tree favoring secondary forest. Trunk has small plank buttresses and light bark. Leaves are simple, alternate, short and rather wide, prominently

toothed, and have two glands (tiny dark bumps) at their base, just above the petiole. The leaf is typically held at an angle to the petiole, a common Alchornea trait. Flowers are minute, on long, thin stalks. Distribution: Known widely but sporadically, never especially common. Found in all forests around the Canal. It appears as good-sized trees in mature forest, but saplings are rare; not typically along roads or in open areas. Recognition: The short, wide leaves are euphorb-like, and with prominent teeth they are easy to recognize. Check for the glands at the leaf base. Beware the remarkably similar Hasseltia floribunda (Salicaceae), which even has glands; Hasseltia has sharper teeth. See also A. latifolia.

Euphorbiaceae

Adelia triloba

Alchornea costaricensis

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Euphorbiaceae Alchornea latifolia

Alchornea latifolia

Alchornea latifolia (achiotillo). A medium-sized

tree of wetter areas. Trunk typically irregular and sometimes fluted; bark brown. Leaves are simple, alternate, only weakly toothed and sometimes not at all. At the base of each leaf are two or four glands

(tiny dark bumps). The tiny flowers are on branched spikes. Distribution: Known widely from c. Panama through Costa Rica in wet and montane forest, including swamps near the Caribbean. Found through most of the Canal Area. It appears occasionally as a good-sized tree in mature forest, and also in secondary forest and sometimes forest edge. Recognition: This species has larger leaves with fewer teeth than A. costaricensis. Check also Salicaceae, especially Hasseltia floribunda. Alchornea contains five more species, all scarce in both Panama and Costa Rica. They all have fairly broad, toothed leaves with glands.

Croton billbergianus

Croton billbergianus

Croton billbergianus (sangrillo, vaquero). A me-

dium-sized, weedy tree. Trunk is cylindrical, straight, light gray. Leaves are large, heart-shaped, light gray below with a thin coat of rough reddish hairs; branchlets have the same rough feel. Older leaves turn orange or yellow, and there are usually a few orange leaves on any good-sized tree. On the petiole just below the leaf there is a pair of glands that are

just tiny, elevated swellings. Breaking a leaf or branchlet produces dripping orange sap. Flowers are small, greenish white, on long and usually terminal stalks. Distribution: A widespread forest-edge species throughout except in the driest zone. In mature forest, only occurs in natural clearings and usually as a shrub or treelet. Abundant along wooded roads in the Canal Area, where it reaches 10 m tall or more; not as common in nonforest areas. Recognition: Such conspicuously heart-shaped leaves are not found on many trees, but two genera in the Urticaceae we do not cover, Urera and Myriocarpa, have very large leaves shaped like those of Croton. The tiny glands help cinch Croton, which is is so abundant near roads in the Canal Area that it is easy to learn quickly. See C. draco, which is quite similar, and also the heart-shaped but toothed leaves of Hura crepitans, also a euphorb.

Euphorbiaceae

Alchornea latifolia

Croton billbergianus

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Euphorbiaceae Croton draco

Croton draco

ramidale (Malvaceae—Bombacoideae). Neither of those species produces red sap though.

Croton draco (sangrillo, sangare, sangre de drago, algodoncillo). Another medium-sized weedy tree

Croton includes 12 more Panamanian species from many parts of the country, especially wet and montane forest. Most have heart-shaped leaves, but a few do not; some have toothed leaves. All produce reddish sap and have glands where the petiole meets the leaf. C. schiedeanus is a very widespread and common species having narrow leaves pointed at both ends (far from heart-shaped); the other species are scarce.

mostly of montane sites. Similar to C. billbergianus, but sap from broken leaves is red, not orange; and leaves have widely spaced secondary veins, can be slightly lobed, and sometimes have fine teeth. There are glands on the petioles just like in other Croton. Flowers are white, on long stalks, conspicuous above the leaves. Distribution: Typically in lower montane forest from c. Panama through Costa Rica, but can be found at low-elevation sites along the Pacific coast too. It is a roadside and edge species like C. billbergianus. Recognition: See C. billbergianus. Leaves are also similar to those of Heliocarpus americanus (Malvaceae—Grewioideae) and the balsa, Ochroma py-

Hevea brasiliensis (rubber, caucho). A tall tree, not native to Panama. Leaves are trifoliate and on long petioles; they often turn orange or yellow while still on the tree. Broken leaves or slashed bark produce white latex—the source of rubber. Distribution: Sometimes found along city streets, and there is a big collection at Summit Garden near Panama City. Occasionally found in the forest, where there were once plantations. Native to Amazonia. Recognition: There are few native species with trifoliate leaves; none have such long petioles and none occur in cities.

Euphorbiaceae

Croton draco

Hevea brasiliensis

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Euphorbiaceae

Hippomane mancinella

Hippomane mancinella

Hippomane mancinella (manzanillo de la pla-

ya). A medium-sized tree of beaches. Trunk has low

branches, and the crown is round and broad. Leaves are small, round at the base, barely toothed, and densely clustered toward the ends of branches The petiole has a small gland near the leaf. Broken leaves or bark produce a milky sap that stings and may cause a rash. Flowers are small, greenish, on long stalks. Fruits are spherical, with hard flesh, and are toxic! Distribution: A coastal species throughout both countries, common and easy to find near mangroves and beaches. Recognition: Easy to spot in its habitat. The leaves resemble those of Hura crepitans, but are smaller and less toothed.

Hura crepitans

Hura crepitans

Hura crepitans (nuno, tronador, havillo, ceibo,

sandbox tree). A forest giant. Trunk can be immense, matching Ceiba (Malvaceae—Bombacoideae) or Anacardium excelsum (Anacardiaceae) in size but without buttresses. Trunks of saplings and mediumsized trees are covered in sharp, cylindrical spines, but these become sparse in giant trees. Leaves are heart-shaped, toothed, with straight and very parallel secondary veins; there is a pair of small, swollen

knobs on the petiole just below the leaf. Broken leaves produce a clear sap that turns milky; it is toxic. Flowers are held on a conical, stalked head, and are deep red with a long corolla when open. Fruit is a small, cubicle capsule, starting green but turning brown. When dry, it explodes and catapults the seed; sometimes the popping detonation is audible from the ground. Distribution: Widespread at low elevation in dry to moist zones, especially along the Pacific slope, but also near the Caribbean in a few areas. Often along rivers in agricultural areas. Recognition: Leaves are quickly learned; note the straight secondary veins. See Croton and other species mentioned there with heart-shaped leaves; all have completely different venation and no teeth. Hura is well known for timber and occasionally grown in plantations. Trees in Bocas del Toro and old forest at Barro Colorado I. are among the largest in Panama.

Euphorbiaceae

Hippomane mancinella

Hura crepitans

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Euphorbiaceae Jatropha curcas

Jatropha curcas

Jatropha curcas (coquillo). A shrub or treelet of farmland and waste areas. The trunk is much branched and has a rounded crown. Leaves are

large, heart-shaped and almost lobed, with three prominent veins arising at the base. Flowers are green, in flat terminal heads; fruits are capsules that split open. Distribution: Mostly in farmland and savannas of the Pacific dry zone. Used in living fencerows in some areas. Recognition: Leaves are similar to those of Croton draco, and see other species mentioned there. All these big-leaved species favor clearings and forest edge; Jatropha is a smaller shrub of the dry zone. A second Jatropha species, also of dry areas, has slightly lobed leaves.

Mabea occidentalis

Mabea occidentalis

Mabea occidentalis (casiquillo). A treelet of mature forest. Several branches emerge together from the small trunk (verticillate branching). Leaves are smooth, simple, alternate, slightly toothed, with undulate margins. The upper side is dark green, and the underside light blue with an iridescent tint, with a fine network of tertiary veins. On young branches, stipules are doubled and angle backward (but they soon fall off). Breaking off a leaf produces a rapid flow of white latex. Small green flowers are in a large, long terminal cluster, as are the small fruit capsules. Distribution: Moist and wet regions throughout, in

the understory of mature forest. Recognition: The bluish leaf undersides are a good character. Several Moraceae (Sorocea and Brosimum) are similar in the tertiary venation and copious white latex, but do not have the light blue color. Virola (Myristicaceae) plus Xylopia and Mosannona (Annonaceae) have bluish leaf undersides, but due to dense hairs, and they have different venation, no teeth, and scarce or no latex. One other Mabea species has very similar, but smaller, leaves, and fuzzy stems. It is mostly known in far nw. Costa Rica.

Manihot esculenta (manioc, yuca). Not illustrat-

ed. A shrub-sized crop native to Amazonia. Has palmately compound leaves like those of marijuana plants. Distribution: Grown in yards and gardens everywhere in Panama and Costa Rica. Recognition: Leaves are reminiscent of those of Pachira (Malvaceae—Bombacoideae).

Euphorbiaceae

Jatropha curcas

Mabea occidentalis

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Euphorbiaceae Pera arborea

Pera arborea

Pera arborea (sapito, clavito, pellejo de gallina, felí). A tall forest tree. Trunk is straight, cylindrical; bark has scaly pores. Leaves are simple, alternate, regularly spaced and held in a flat plane, dark and

smooth green above, light green and slightly scaly below. Flowers are small, white, with yellow stamens, born directly off the branches. Fruits are capsules. Distribution: Found widely from dry to wet zones in both countries, and sometimes in lower mountains. Recognition: Very nondescript and difficult to learn, with leaves much like those of several other nondescript species. Quite similar to Amanoa and Margaritaria (Phyllanthaceae) and also Drypetes (Putranjivaceae) and Bourreria (Boraginaceae). But Amanoa and Bourreria have distinctive trunk characters. Pera has two more species in Panama, both rare. One of them has opposite leaves.

Sapium glandulosum

Sapium glandulosum

Sapium glandulosum (olivo). A tall tree of forest

edge. Trunk is straight, cylindrical, with small plank buttresses at the base; bark has long vertical fissures dividing regular, rectangular plates. Leaves are shiny green, finely toothed, and held at various angles from the branchlets. There is a pair of stalked glands on the petiole where it meets the leaf. Broken leaves gush white latex. Flowers are minute, pink, on long, thin terminal stalks. Fruits are capsules with six sections. Distribution: Widespread from dry to wet zones throughout. Often forest edge, sometimes

roadsides, and in rural areas used in fencerows. But also in mature forest, generally in natural clearings. Recognition: Easy to recognize from the alligator bark, small buttresses, and distinctive leaves, even without checking for abundant latex. Might be confused with Ficus trees (Moraceae) or the ornamental frangipani (Plumeria [Apocynaceae]); both have similar shiny leaves and copious white latex, but lack glands and teeth and have different bark. Tetrorchidium gorgonae has similar glands but is otherwise different. Three other Sapium species are reported in Panama. Two are montane, the third is known from lowlands in much of Costa Rica but is very rare in Panama. Moreover, we have observed trees resembling S. gladulosum yet not identifiable to one of the known species, and so likely another species. All have abundant white latex and glands near the leaf base, though in most the glands are not stalked.

Euphorbiaceae

Pera arborea

Sapium glandulosum

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Euphorbiaceae

Tetrorchidium euryphyllum

Tetrorchidium euryphyllum

Tetrorchidium euryphyllum. A cloud-forest tree. Trunk often has low branches. Twigs are square and hollow. Leaves are simple, alternate, somewhat bunched toward the branch ends, rounded and fairly

wide, with nearly blunt tips and few widely spaced secondary veins. Leaf underside is much lighter than the upper side. There is a pair of small, dark spots at the base of the leaf where it meets the petiole. Latex drips from broken parts. Flowers are tiny and white; fruits are capsules. Distribution: On scattered low mountains from c. Panama through Costa Rica. Recognition: Might be confused with Terminalia in the Combretaceae. Hieronyma oblonga (Phyllanthaceae) is reminiscent in leaf shape, but has brownish scales on the leaf underside. The small spots on the leaf base are subtle and resemble glands of Alchornea (another euphorb) or Hasseltia (Salicaceae), but the latter both have toothed leaves.

Tetrorchidium gorgonae

Tetrorchidium gorgonae

Tetrorchidium gorgonae. A wet-forest tree.

Trunk often has low branches. Twigs are square and hollow. Leaves are simple, alternate, somewhat bunched toward the branch ends, rounded and fairly wide, with blunt tips, and a few widely spaced secondary veins. Old leaves turn yellow on the tree. Broken parts leak small amounts of white latex. On the petiole just below the leaf are two tiny green stalks. Flowers are minute. Distribution: Known from widely scattered sites in lower montane and Caribbean wet forest. Fairly common in secondary forests and along roads of the Caribbean slope near the Canal. Recognition: See similar species mentioned un-

der T. euryphyllum. The little stalked glands of this species are an excellent character; they are like those of Sapium, but Sapium has much different leaves and more latex. Three more species of Tetrorchidium are found in Panama. Two are very seldom seen. The third, T. costaricense, is from mountains and mostly known in Costa Rica; it also has glands and fairly wide leaves. Fifteen genera remain in this big family: Acidoton, Adenophaedra, Alchorneopsis, Caryodendron, Cleidion, Cnidoscolus, Conceveiba, Euphorbia, Garcia, Gymnanthes, Maprounea, Pausandra, Sagotia, Senefeldera, and Tacarcuna. Three of those have two species, and the rest just one. Most are very seldom seen, but those that are fairly well known include Acidoton nicaraguensis, a wet-forest treelet with toothed leaves much like those of Acalypha; Cnidoscolus urens, a small, weedy dry-zone tree having wide, lobed leaves; Euphorbia elata, a wet-forest treelet with long, narrow leaves held erect at the top of the stem; and Pausandra trianae, a small wet-forest tree with long, toothed leaves broadest near the tip.

Euphorbiaceae

Tetrorchidium euryphyllum

Tetrorchidium gorgonae

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Fabaceae

Fabaceae

Brownea macrophylla

Brownea macrophylla

These are the legumes, the family of peas, beans, clover, rosewood, mesquite, and many other wellknown plants of all types. The family is common and widespread throughout the world, often predominant in deserts and common in temperate grasslands, and also one of the world’s major groups of tropical trees. There are 18,000 species, ranking it as the third largest plant family (after the sunflowers [Asteraceae] and the orchids [Orchidaceae]); 6,700 are known in the American tropics. Many of those are small herbs or lianas, but there are large numbers of tree species, including very tall trees. Indeed, it is the second largest family in the Panama tree checklist, with 234 species (barely behind the Rubiaceae, which has 236). With a couple minor exceptions, trees in the Fabaceae have compound leaves. Many have rounded leaflets, and many have small or tiny leaflets. Some have bipinnate leaves, meaning leaves made up of leaflets, and leaflets in turn subdivided into subleaflets. The jargon for a subleaflet is a pinnule, and we will use the words leaflet and pinnule frequently and carefully. Bipinnate leaves are usually obvious, and few non-Fabaceae have them. Even more distinct are the petioles: the base of each, where it meets the branch, is cylindrical, thickened, and green. Likewise, the petiolule (leaflet stalk) often has a similar swelling. There are very few species in the family lacking the swollen petiole base, and conversely, the character is very rare outside the family. It is the fruit, however, that provides the best known character of the family: the seed pod. Most species have pods, but there are exceptions having winged seeds like those of the North American maples (the winged seeds are flattened seed pods having one seed); in addition there are a few Fabaceae with fleshy fruits, lacking a pod. Flowers are also distinctive, but we describe them under subfamilies. Because of the large number of species, the Caesalpinia coriaria

Caesalpinia coriaria

Fabaceae are usually treated in three subfamilies, Caesalpinioideae, Mimosoideae, and Papilionoideae, and we follow that approach here. This is a useful division, because the three subfamilies are usually distinct; nevertheless, without flowers, there are quite a few species that do not obviously fall in one of the three groups. Remember, though, that all three groups share the core Fabaceae traits—compound leaves with a characteristic petiole base—and you will do fine just recognizing the family and not worrying about the subfamilies. Indeed, despite the great number of species, Fabaceae are fairly easy to learn: most species are readily identified to the family, and many species within the family are distinct and easily recognized. The challenge is simply the very large number of species.

Caesalpinioideae (subfamily within Fabaceae). This is the smallest of the three sub-

families, with 2000 species worldwide and 1200 in the American tropics; most are trees or shrubs. Caesalpinioid leaves are usually even-pinnate, with no terminal leaflet, and leaflets are opposite one another. Neither character holds for all, though, and some caesalpinioids have bipinnate leaves and tiny leaflets, like the mimosoids. Caesalpinioid flowers are regular, with only slight differences in size among the five petals (the lower petal may be slightly enlarged); they are often large and yellow or pink. If you come across a tree that is clearly a legume and has an even number of leaflets, it is either a caesalpinioid or the genus Inga in the Mimosoideae. Panama has 54 species of trees and shrubs in the Caesalpinioideae, of which we illustrate 10 and describe 6 more.

Brownea macrophylla (rosa de monte, cuchilli-

to). A wet-forest treelet. Leaves are very long, evenpinnate, with many pointed leaflets. Young leaflets are pink and droop downward, creating a conspicuous and highly distinctive display. Flowers appear on the trunk in big, brilliant red bunches. Fruits are narrow, woody, brown pods. d istribution: Only in the eastern half of Panama, in lowland wet forests. In the Canal Area, found in swampy areas near Madden Dam. Recognition: Possibly confused with Myrtaceae if the long leaves are not recognized as compound. The thickened, cylindrical petiole base is readily evident however. Guarea guidonia (Meliaceae) also has long leaves with many leaflets, and Inga nobilis (Mimosoideae) has very similar leaflets, but fewer. Flowers of Brownea are unmistakable. Brownea has a second species, B. rosa-de-monte, used as an ornamental; its natural range is not known. The leaves are similar to those of B. macrophylla, but flowers are orange.

Caesalpinia coriaria (agallo). A medium-sized tree of dry zones. Leaves are bipinnate, with many

Fabaceae—Caesalpinioideae

177

Brownea macrophylla

Caesalpinia coriaria

tiny, blunt leaflets having small black spots below. Flowers are whitish with yellow stamens. Fruits are long pods twisted in on themselves. d istribution: Only occurs in the driest zones of c. Panama and nw. Costa Rica, in pastures and scrubland. Recognition: See especially Mimosa in the Mimosoideae, but it has spines; Parkinsonia is another caesalpinioid in dry areas with tiny leaflets, but it is quite different. The tiny leaflets of C. coriaria resemble those of Enterolobium cyclocarpum (Mimosoideae),

which grows in similar habitat but is a much bigger tree with a different fruit; juveniles of the two, which are seldom seen, are similar, but look for black spots on the leaflets of this species and tiny glands on the petiole in Enterolobium. A second native species of Caesalpinia grows in the dry zone, and has similar bipinnate leaves. C. pulcherrima is a small, orange-flowered shrub planted in gardens, introduced from the West Indies.

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Fabaceae—Caesalpinioideae Cassia moschata

Cassia moschata

Cassia fistula (caña fistula). Not illustrated. A small tree native to India. Has six to eight shiny green leaflets, far fewer than C. moschata. Bright yellow flowers dangle downward under leaves and are often present. d istribution: Planted in cities and seen all over Panama City. Recognition: Never seen outside towns so not difficult to learn.

Cassia moschata (casia amarilla, carao, caña fístula). A medium-sized tree of secondary forest. Leaves are even-pinnate, with many small leaflets that are round at the tip. Flowers are orange, produced in great numbers toward the end of the dry season. Fruits are long, pencil-shaped, brown pods. d istribution: Abundant along roadsides and in secondary forest near Panama City. Barely known in Costa Rica or elsewhere in Panama. Recognition: Usually known from the flowers, visible at great distance. Leaves are typical of the genus Cassia and closely related Senna. Abarema (Mimosoideae) has similar leaflets, but leaves are bipinnate; this is not at all obvious in big trees, but Abarema is a rare forest tree. Another Cassia species, C. grandis, is native to Panama. It has leaves like those of C. moschata, but flowers are pink; it is much less common.

Copaifera aromatica

Copaifera aromatica

Copaifera aromatica (cabimo). A splendid tall tree. Trunk straight and often unbranched until very high. Bark yellowish and smooth, with peeling sheets. Has fragrant, resinous sap, exuded from cut bark. Leaves are even-pinnate, with four or six leaf-

lets; the basal pair is alternate whereas the terminal pair is opposite. Leaflets are symmetric, and have tiny translucent spots, visible with magnification only. Flowers are small and white; fruits are round pods holding large seeds. d istribution: In Panama, known from the dry zone near the Pacific, especially along streams, seldom in open areas. There are only a few widely scattered records from Costa Rica. Recognition: Not many even-pinnate species have only four or six leaflets. See Prioria, which always has four leaflets. Crudia has similar leaflets, but they are asymmetric at the base. A second species of Copaifera is widely scattered but rare in Panama.

Fabaceae—Caesalpinioideae

Cassia moschata

Copaifera aromatica

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Fabaceae—Caesalpinioideae Crudia acuminata

Crudia acuminata

Crudia acuminata (algarrobillo). A large tree of

dry zones. The trunk is big but often has low branch-

es. Leaves are odd-pinnate, and the leaflet pairs are alternate along the rachis; the leaflet base is asymmetric. White flowers are on spikes; fruits are short, round pods. d istribution: Known for certain in Panama only in one area of the Pacific slope. It is common there along big rivers (a specimen from high mountains is a misidentification). Rarely seen in Costa Rica. Recognition: In the same habitat, Copaifera is even-pinnate and Hymenaea has just two leaflets. There are some papilionoids with similar leaves (Acosmium, Dalbergia, and Pterocarpus), but they do not occur in dry areas.

Cynometra bauhiniifolia

Cynometra bauhiniifolia

Cynometra bauhiniifolia (pezuña de vaca). Not illustrated. A large but very rare forest tree. Has two leaflets, each with a tiny notch at the tip. Flowers are white. Fruits are not pods, but fleshy with a large seed. d istribution: Very rare in c. and e. Panama, usually along rocky streams. Recognition: Easy to confuse with Hymenaea or Peltogyne, but the tiny notch is distinguishing.

Hymenaea courbaril

Hymenaea courbaril

Hymenaea courbaril (algarrobo, guapinol). A tall tree of drier areas. Trunk is straight, unbuttressed, usually cylindrical but sometimes slightly fluted or irregular. In the forest, it can be a magnificent, large tree, but it is smaller in farmland. Leaves have two asymmetric leaflets; they look like one large leaf that has been torn down the middle. There are tiny trans-

lucent spots in the leaflets. Flowers are mediumsized, white, and in dense clusters. Pods are cinnamon-colored ellipsoids. d istribution: A tree of the drier half of the isthmus throughout, including the very driest zone. Frequently in riverside forest, also farmland. Occasionally in wetter areas and in mature forest. Recognition: Leaflets in pairs are unusual and distinctive, and should be recognizable even high in the canopy, at least with binoculars. A few other species have them, including Cynometra and Peltogyne, neither of which we illustrate. See also Pithecellobium unguis-cati (Mimosoideae), which has similar leaves, though bipinnate, and also has spines. Hymenaea is a well-known tree in rural areas, considered useful for fruit and wood, and bark resins are used in various ways and have been thoroughly studied chemically.

Fabaceae—Caesalpinioideae

Crudia acuminata

Hymenaea courbaril

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Fabaceae—Caesalpinioideae Parkinsonia aculeata

Parkinsonia aculeata

Parkinsonia aculeata (árbol sarigua, palo verde). A small tree of Panama’s driest zone. Trunk and

branches have small, sharp thorns. Leaves are bipinnate, but this is not obvious because the first division includes just two long rachises; each has dozens of tiny pinnules that often fall off. Most of what you see are green leaf rachises, which do most of the photosynthesis. Fruits are small, slender pods with swellings where the seeds are. d istribution: Only in a few dry sites in both Panama and Costa Rica. Recognition: There are no other trees like this in the area, with tiny, sparse leaflets. Relatives in the deserts of Mexico and the sw. United States are called palo verde.

Peltogyne purpurea

Peltogyne purpurea

A second species of Peltogyne, P. mexicana, is known in the Darien.

Peltophorum pterocarpum (acacia africana).

Peltogyne purpurea (nazareno, purple heart).

Not illustrated. A tall tree of wet areas. Leaves have two leaflets. d istribution: Only in the Darien and the Osa Peninsula. Recognition: Much like Hymenaea. The tree gets its name from the color of its heartwood, and it yields good timber for furniture and floors.

Not illustrated. An Asian ornamental. Leaves are bipinnate, with small, rounded pinnules. It is covered in yellow flowers through the dry season, each showing a bit of orange (the stamens) from a distance. Fruits are small, flat pods that typically stick upward above the leaves. d istribution: Planted in many urban areas and common along Panama’s streets. Most likely to be confused with Cassia moschata, which is also abundant around Panama City, but Cassia has once-pinnate leaves and orange flowers.

Prioria copaifera

Prioria copaifera

Prioria copaifera (cativo). A tall forest tree.

Trunk is straight, unbuttressed, and generally unbranched until well above the ground, to nearly 50 m tall and thus one of the tallest trees in Panama. The crown is deep, with dense foliage. Leaves have exactly four leaflets, shiny on the top and asymmetric. Flowers are small, white, on many long stalks. Fruits are flat, brown, circular pods, concave on one side, with a single large seed, often seen on the ground beneath big trees. d istribution: Best

known in swamp forests in the Darien, where it forms large stands called cativales just above the mangroves. Also found along both coasts north through Costa Rica, usually in swamps, though also in upland forest. In the Canal Area it is very common in upland sites, for example in Barro Colorado I. old growth. Numerous both as a sapling in the shade and as a canopy adult; never in open areas. Recognition: Easy to recognize as a sapling or small tree from the four leaflets. The only other nonliana in the area with four and only four leaflets is Senna dariensis, which is a small, leaning shrub. See also Copaifera. As a large tree, leaves are difficult to see, and Prioria is then not easy to identify. Check for leaves on the ground, because the asymmetric leaflets are distinctive, and also look for the woody fruits. Cativo is one of the most commonly harvested timber trees of Panama, because of its large size and its tendency to form nearly monospecific stands in swamps along rivers.

Fabaceae—Caesalpinioideae

Parkinsonia aculeata

Prioria copaifera

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Fabaceae—Caesalpinioideae Schizolobium parahyba

Schizolobium parahyba

Schizolobium parahyba (indio, tinecú, cigarrillo, gallinazo, cucharo). A tall tree of secondary

forest. Trunk is straight, with light-colored bark and small cinnamon-brown buttresses. The crown is wide but open. Leaves are long, bipinnate, with many small, rounded pinnules. Juveniles lack branches, having a single straight stem with huge

leaves emerging just at the top and thus resembling tree ferns in form. Flowers in December (near the Canal), and big trees are conspicuous at great distance when covered in the yellow flowers. The pod is flattened, brown, round at one end and pointed at the other. d istribution: Very common around the Canal and along Pipeline Rd. in Soberania; seldom seen elsewhere, but known widely. Recognition: The bipinnate leaves resemble those of Jacaranda (Bignoniaceae), and juveniles of the two species have the same growth form, but Jacaranda has pointed leaflets and opposite leaves. The brownish buttresses of Schizolobium are a good character in bigger trees. When flowering, it can be confused with Tabebuia guayacan (Bignoniaceae), but Tabebuia does not flower until February.

Senna dariensis

Senna dariensis

Senna dariensis (ictericia, barbaso). Not illustrated. A straggling shrub of forest edge. Leaves always have four leaflets, the end pair asymmetric. Leaf underside is light green, almost blue. The flowers are large, in conspicuous clusters, light yellow, and the pod is slender, cylindrical, like a pencil. d istribution: In natural forest gaps, forest edge, along roads in most of Panama, but not in Costa Rica. Recognition: Leaflets are held in the same pattern as in Prioria, but this will always be distinguished by its straggling form.

Senna reticulata

Senna reticulata

Senna reticulata (laureño). A small tree of open

areas. Leaves are long and have many round-tipped leaflets. Flowers are often present, in conspicuous yellow clusters above the leaves. The pod is long and flattened, with crosswise ribs. d istribution: Grows

along lakeshores, swamps, waste areas, on both slopes in Panama to c. Costa Rica. Recognition: Several Cassia have similar leaves, plus note the large number of Senna species. S. reticulata is best known as the one growing in open areas and often near water. Eighteen other species of Senna are known in Panama. They are shrubs to treelets with four to many small to medium-sized leaflets and yellow flowers. S. papillosa is a fairly common species in w. Panama and Costa Rica, resembling S. dariensis. A couple others are fairly common, but many are seldom collected and poorly known.

Fabaceae—Caesalpinioideae

Schizolobium parahyba

Senna reticulata

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Fabaceae—Mimosoideae Tachigali versicolor

Tachigali versicolor

Tachigali versicolor (suicide tree). A tall forest

tree with large plank buttresses, unbranched until near the top. The bark is smooth, reddish brown; the tops of the buttresses are especially reddish. Saplings have a ribbed, reddish stem, and small branches on big trees are ribbed in the same way. Leaves are long, even-pinnate, with 14–16 pointed leaflets; basal leaflets are smaller. The leaf rachis is reddish with dense hairs, and leaf veins are reddish. At the growing tip of each stem is a bizarre stipule having the form of a miniature leaf—with many miniscule ‘leaflets’. Tachigali takes a remarkable approach to reproducing. Trees grow huge before they finally reproduce for the first time, producing light brown flowers in clusters above the crown that develop over a whole year into flattened pods with one seed each. Then the giant tree drops its leaves, sheds the winged seeds into the wind, and dies. d istribution: Very common in the Canal area from moist to wet sites; otherwise known Abarema barbouriana

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MiMosoideae (subfamily within Fabaceae). Another of the three subfamilies of Fabaceae,

and the second largest, with 3200 species worldwide and 1400 in the American tropics. Panama has 100 species of trees and shrubs in the subfamily, of which we illustrate 24 and describe 12 more. Mimosoids typically have bipinnate leaves with tiny, narrow leaflets, so leaves resemble feathers; however, the biggest genus, Inga, is not bipinnate and has larger leaflets. Most have tiny glands on the petiole or leaf rachis. Members of this subfamily Acacia melanoceras

only at a couple sites in Costa Rica. Where found, saplings in mature-forest understory are numerous. Never seen in open areas. Recognition: The pointed leaflets with reddish veins do not immediately suggest Fabaceae, but the cylindrical petiole base does. Saplings are easy to recognize from their ribbed, reddish stems and peculiar stipules at the top. As an adult, the buttresses and smooth, reddish-brown bark are a good characters but not diagnostic without seeing the compound leaves. Compare two Sapindaceae: Talisia nervosa has compound leaves and a stipule just like Tachigali; Cupania scrobiculata has compound leaves and a ribbed stem, but toothed leaflets. There are five more caesalpinioid genera: Bauhinia, with seven species, and Browneopsis, Dialium, Mora, and Sclerolobium, each with a single species. Bauhinia is distinctive, with two leaflets connected at the base, as if they are in the process of splitting apart; it is best known as a widespread liana called monkey ladder, but several are small trees. Dialium has only a few pointed leaflets, and they are alternate, unlike most Caesalpinioideae; it is a tall wet-forest tree. Sclerolobium resembles Tachigali and is seldom collected. Mora has four leaflets, like Prioria, but they are symmetric; like Prioria, it grows in swamps near mangroves, but it is only known in far e. Panama and the Osa Peninsula.

have a distinct flower, different from that of the other Fabaceae: petals are small and not conspicuous, and numerous long stamens extend well beyond the petals. They are like little balls of stamens, resembling small brushes or cheerleader’s pom-poms.

Abarema barbouriana. Not illustrated. A tall forest tree. Twigs have fine and dense reddish hairs. Leaves are bipinnate, and pinnules are small, rounded. There is one small, circular gland between each pair of leaflets. Flowers are white balls of stamens. Pods are flattened, curved. d istribution: In mature forest around the Canal, not very common. Elsewhere very rarely known. Recognition: Numerous leaflets are small, but not tiny as in the featheryleaved Enterolobium or Acacia. A second species of Abarema, A. macradenia, is similar except for having just one large disk-shaped gland on each leaf, between the lowest pair of leaflets. It is rare.

Acacia melanoceras

Acacia melanoceras (cachito, cuernito, bull

thorn acacia). A small forest tree with ant thorns. Branches have big, swollen thorns, always in pairs and thus resembling bull horns. If you make the mistake of breaking a thorn, ants come out to defend their tree, and they bite hard. Leaves are feathery bipinnate. Flowers are small, yellow fuzzballs in dense

Fabaceae—Mimosoideae

187

Tachigali versicolor

Acacia melanoceras

heads. Pods are slender, cylindrical, pointed at one end. d istribution: Only in c. Panama, where found in dry and wet areas, in mature forest shade but also open areas, always sparsely. Where they are found, it is nearly always as a small group of trees within a few meters of each other. Recognition: Unmistakable thorns, but note that other Acacia have these.

Panama has nine more Acacia species as trees, plus more as lianas. All have bipinnate leaves with tiny leaflets, and some others have thick bull thorns. Most are in the dry zone, and better known in Costa Rica. Acacia is one of the largest genera of trees in the world (especially Australia and Africa), but it is not important in tropical rainforest.

188

Fabaceae—Mimosoideae Albizia adinocephala

Albizia adinocephala

Albizia adinocephala. Not illustrated. A slender

roadside tree. Leaves are bipinnate, with pointed, medium-sized leaflets, smaller than Inga but larger than the feathery-leaved mimosoids. Flowers are solitary white fuzzballs; pods are long and flattened, ribbed to indicate where seeds are. d istribution: In

the dry zones of c. Panama and nw. Costa Rica, where seen in agricultural areas. Very common along roads on the Pacific side of the Canal Area. Recognition: Much larger leaflets than most of the bipinnate mimosoids with their feathery leaves. Might look like Inga, but Inga is not bipinnate and has leaf glands. Fairly easy to learn because it is so common where it occurs. Three other Albizia species are native to Panama. A. niopoides is a dry-zone species with tiny leaflets, like those of Enterolobium or Acacia. Others have larger leaflets like those of A. adinocephala and are very rare. There are also two exotic species planted as ornamentals.

Calliandra magdalenae

Calliandra magdalenae

Calliandra magdalenae (gallito). Not illustrated. A familiar garden tree. Leaflets are medium-sized, asymmetric. Flowers are purple fuzzballs, nearly always present, attractive to hummingbirds. Pods are long, flattened, with raised lips running the length of Cojoba rufescens

Cojoba rufescens

Cojoba rufescens (coralillo, harino). A small tree

of all habitats. The trunk is often leaning and branched near the ground. Leaves are even-pinnate, with six to eight leaflets. Sides of leaflets are wavy, and leaflets near the base are reduced in size. There is a tiny circular gland on the leaf rachis between each pair of leaflets. Flowers are small, in dense globEnterolobium cyclocarpum

Enterolobium cyclocarpum

either edge. d istribution: An ornamental in cities and farms of the Canal Area, but native and reported along the Pacific slope in a variety of habitats. Never inside the forest. Recognition: Generally seen in yards, where it is readily learned from its purple flowers. Calliandra has eight native species in Panama. C. laxa is found in wet forests along the Caribbean coast near the Canal. Other species are rare. Leaves are variable, some with many small leaflets, and others with just two or four larger leaflets. They all have fuzzballs for flowers, with very long extruding stamens, varying from white to pink; most have fruits with raised edges.

ular heads. The pods are red and twisted, and they split open to reveal shiny black seeds. d istribution: Widespread from Panama to e. Costa Rica; mainly in drier areas along the Pacific slope, but also at wet sites. Abundant and easy to see nearly everywhere in the Canal Area. Often at forest edge or lake margins, not inside the forest. Recognition: Resembles the genus Inga in leaflet form and the gland between each pair. C. rufescens should be readily recognized, though, by the undulate leaflets, a consistent trait not found in Inga, and the distinctive fruits it bears. Two other species of Cojoba are rare in mountains of w. Panama, and more widely known in wet lowlands of Costa Rica. One has much smaller leaflets, the other larger; both have rachis glands and fruits like those of C. rufescens.

Enterolobium cyclocarpum (corotú, guanacas-

te, ear tree). A huge pasture tree. The trunk has small buttresses that extend into roots that can be traced above the ground for great distances. It often forks near the ground, and big branches are near horizontal and sometimes touch the ground. The crown is usually very broad and can be wider than it is tall. In contrast to this typical pasture appearance, trees will sometimes grow a straight, tall trunk inside the forest. Leaves are bipinnate, with tiny, feathery leaflets. On

Fabaceae—Mimosoideae

189

Cojoba rufescens

Enterolobium cyclocarpum the leaf rachis, between each pair of leaflets, there is a tiny raised dishlike gland (as in Inga). The flowers are small balls of stamens, cream-colored, and single trees smell sweet from afar when covered with flowers. The pods are green at first, but mature brown, and are shaped like an ear. Cattle, goats, and horses eat the fruits and disperse the seeds. d istribution: A dominant giant of pastures and grasslands of the Pacific slope throughout, and a very common component of secondary forest regrowing from pasture. Saplings are never in the forest; this species only germinates successfully in open fields where there are

livestock around. Recognition: This and the other Enterolobium look very similar and are easily confused, and both are abundant in secondary forests around the Canal, but E. cyclocarpum is the predominant pasture species. Leaflet size also distinguishes them: leaflets of E. cyclocarpum are about twice as large (so whereas leaflets of E. cyclocarpum are tiny, those of E. schomburgkii are miniscule threads). There are other legumes with fine, compound leaves (see Acacia, Mimosa, and Prosopis, plus Caesalpinia and Parkinsonia in the Caesalpinioideae), best distinguished by tree form, size, and habitat.

190

Fabaceae—Mimosoideae Enterolobium schomburgkii

Enterolobium schomburgkii

Enterolobium schomburgkii (corotú de montaña, dormilón, harino, guábilo, zarza, jarino). Very similar to the previous species, but leaflets are even tinier, barely 1 mm wide, almost threads. This species never reaches the large size of E. cyclocarpum, and has smaller fruits. d istribution: Usually seen in secondary forest, not common in pasture, and mostly known on the Pacific slope of the Canal Area; rare in Costa Rica. The forest on the hill above Gamboa is dominated by E. schomburgkii. Recognition: See E. cyclocarpum. No other tree around has such tiny leaflets.

Inga bella

Inga bella

Genus Inga. One of the largest tree genera in

tropical America. With 55 species in Panama, it ranks second after Miconia in tree species richness. Despite so many species, it is easy to recognize as a genus: all species have even-pinnate leaves, and there are small, circular glands on the leaf rachis (the stalk carrying the leaflets) between each pair of leaflets. The only trees in Panama that have glands between the leaflets and are not Inga are other mimosoids: Abarema, Cojoba, and Pithecellobium. Nearly all Inga are medium-sized trees, never tall

(we will note cases where this does not hold). Most are in the forest, though some appear commonly along wooded roadsides. Remarkably, given the high diversity, most species of Inga are not difficult to learn, because they differ from one another in the number of leaflets and in whether the rachis is winged. Flowers of Inga are white pom-poms, typical of mimosoids, and pods are typically long, flattened, with ridged edges, often curved, and hold several seeds (where otherwise, we indicate). The pods of some species have soft, white, sweet, edible pulp. Local names for all Inga are guaba, guabo, guabito, or some rendition thereof.

Inga bella. Small as Inga go, a treelet or small tree. Four leaflets, with the outer pair much larger. Rachis is winged. d istribution: Sparsely known, in wet forests in the Canal Area and the Osa Peninsula. Recognition: I. chocoensis and I. spectabilis also have four leaflets and a winged rachis, but the former has red hairs and the latter has larger, wrinkled leaflets.

Fabaceae—Mimosoideae

Enterolobium schomburgkii

Inga bella

191

192

Fabaceae—Mimosoideae Inga chocoensis

Inga chocoensis

Inga chocoensis. Leaves are similar to those of I.

bella, but with a dense coat of red hairs on veins and branchlets. Sometimes they have an extra pair of glands along the leaflet stalks (in addition to the main gland on the rachis). d istribution: Cloud forests and wet forests, sparsely from the Canal through Costa Rica. Common at Cerro Campana. Recognition: I goldmanii has similar red pubescence, but more leaflets. See I. bella and I. spectabilis, both of which lack red hairs.

Inga cocleensis

Inga cocleensis

Inga cocleensis. Usually with six or seven pairs of leaflets, sometimes four or eight. No wing on the ra-

chis. Basal leaflets are only slightly smaller than the outer. Twigs and leaf rachis have fine red hairs. Fruit is not a flattened pod, but rather long, slender, and cylindrical, with ribs. d istribution: Widely known in c. and w. Panama especially on the Pacific coast, but also sparsely on the Caribbean coast, including some records in Costa Rica. Often common along rivers and wooded roadsides. Recognition: I. thibaudiana has very similar leaves, but its glands are taller and the fruit is curved; I. multijuga is also similar.

Fabaceae—Mimosoideae

Inga chocoensis

Inga cocleensis

193

194

Fabaceae—Mimosoideae Inga edulis

Inga edulis

Inga edulis. Not illustrated. Native to South America, but cultivated in Panama and Costa Rica. Leaves have 8–10 leaflets and a winged rachis. The fruit is slender, cylindrical, and absurdly long, well over 1 m. d istribution: Common along roads in Bocas del Toro; occasionally seen elsewhere in farmland of both countries. Recognition: See pictures of I. oerstediana, which has similar leaves.

Inga laurina

Inga goldmanii

Inga goldmanii. Leaves with three to five leaflet pairs. The outer pair of leaflets has three glands between them, one on the rachis and one on each of

the leaflet stalks. Veins are prominent, so the leaves look rugose or wrinkled. The rachis is prominently winged between the leaflets. Young leaves, petioles, and branchlets have a dense coat of hairs. d istribution: Central Panama through Costa Rica. Common in the Canal Area, where it is found in most forests but never common; sometimes along roads too. Recognition: Three glands at the base of the outer leaflets make a good character. Only I. chocoensis has three glands like this, and it has only four leaflets. I. mucuna is another hairy Inga, but does not have glands in threes. I. goldmanii has larger flowers than most Inga.

Inga goldmanii

Inga laurina

Inga laurina. Leaflets in two pairs, with the outer pair slightly larger than the inner. The base of each leaflet is slightly asymmetric, and the leaflets are shiny green, without hairs. The rachis has very nar-

row green ridges on each side. d istribution: Pacific coast of Panama and sparsely on the Caribbean near the Canal and in Costa Rica. Found in disturbed secondary forest as well as old growth; not especially common. With I. cocleensis and I. vera, one of the few Inga species occurring in drier areas. Recognition: The closest Inga is I. punctata, which has four to six leaflet pairs, but leaflets of I. punctata are larger and lack the shiny luster of I. laurina. I. marginata has similar leaflets but a winged rachis. See also Zygia, which has glands and leaflets like those of I. laurnia, but look closely: leaves of Zygia are bipinnate, unlike Inga.

Fabaceae—Mimosoideae

Inga goldmanii

Inga laurina

195

196

Fabaceae—Mimosoideae Inga marginata

Inga marginata

Inga marginata. Four leaflets per leaf, with the

outer pair slightly larger than the inner. Leaflets are shiny green, without hairs. Rachis winged. d istribution: Throughout Panama and Costa Rica, from moist to wet zones. Recognition: I. laurina has four shiny leaflets, but lacks the winged rachis. See also I. umbellifera and Zygia.

Inga mucuna

Inga mucuna

Inga mucuna. Usually three pairs of rather large leaflets, a winged rachis, and dense red pubescence. d istribution: Central Panama only, where usually in secondary forest. Recognition: Much like I. goldmanii, but lacking the extra pair of leaf glands.

Inga multijuga

Inga multijuga. Not illustrated. Leaflets in four or five pairs, with basal pairs only slightly reduced relative to outer pairs. d istribution: In wet lowlands of both countries; widespread from moist to wet in c. Panama. Recognition: See photo of I. thibaudiana, which is very similar.

Inga multijuga

Fabaceae—Mimosoideae

Inga marginata

Inga mucuna

197

198

Fabaceae—Mimosoideae Inga nobilis

Inga nobilis

Inga nobilis. Narrow leaflets in three or four pairs.

The basal pairs are smaller. Veins beneath are reddish due to fine hairs, especially on younger leaves. Rachis is not winged. d istribution: Central Panama and very sparsely in Costa Rica. Often in secondary forest and roadsides near the Canal. Recognition: I. ruiziana is similar but has four or five leaflet pairs. Cojoba rufescens has similar leaflets but with undulate margins. Guarea guidonia (Meliaceae) might be confused with I. nobilis or I. ruiziana at a glance, but it lacks rachis glands.

Inga oerstediana

Inga oerstediana

Inga oerstediana. Leaflets in three to five pairs.

Rachis is winged. Leaf veins, rachis, and twigs are reddish due to fine pubescence. Fruits are cylindrical. d istribution: Principally a lower montane species in Panama and Costa Rica, but also in lowland wet forest. Recognition: See I. edulis, which is sometimes considered the same species.

Fabaceae—Mimosoideae

Inga nobilis

Inga oerstediana

199

200

Fabaceae—Mimosoideae Inga pezizifera

Inga pezizifera. Leaflets in four or five pairs. Ra-

Inga pezizifera

chis has narrow wings, sometimes barely visible. d istribution: Wet lowlands, lower montane, and moist forest. Common and widespread near the Canal. Recognition: I. nobilis is similar, but entirely lacks wings; I. pezizifera may sometimes appear wingless. See also I. ruiziana.

Inga punctata

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outer pair much larger. Rachis not winged. d istribution: Found throughout Panama and Costa Rica in lowland moist to wet, and lower montane forest, on both the Pacific and Caribbean slopes. The most commonly seen Inga in Panama. Recognition: Like I. laurina, but leaflets are not smooth and shiny as in that species.

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Inga ruiziana

Inga ruiziana

Inga ruiziana. Not illustrated. Leaflets in four or

five pairs, with basal pairs smaller. Leaf rachis often has very narrow and incomplete wings. Young leaves are pubescent, but older leaves not. d istribution: Widespread in wet to moist lowlands. In Panama, seemingly always rare. Recognition: Like I. pezizifera, with a partial wing. See also I. laurina, which is sometimes considered the same species.

Fabaceae—Mimosoideae

Inga pezizifera

Inga punctata

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Fabaceae—Mimosoideae Inga sapindoides

Inga sapindoides

Inga sapindoides. Leaflets in three pairs; not

much size difference among them. Rachis has a narrow wing. Leaflets, rachis, and twigs have fine, dense, yellowish or reddish hairs. d istribution: Widespread in lowlands of both countries, from Pacific to Caribbean and even some lower montane sites. Recognition: Most like I. goldmanii, which has similar-sized leaflets, but I. goldmanii has denser pubescence and three glands at the base of each leaflet pair. Other pubescent species have smaller leaflets, whereas other species with the winged rachis do not have pubescence.

Inga spectabilis

Inga spectabilis

Inga spectabilis. Leaflets in two pairs, the outer

pair very large. Veins are impressed, giving leaf a rugose appearance. Rachis is winged. Branchlets are square in cross section (try rolling between your fingers). Fruit is cylindrical but constricted between seeds. d istribution: Everywhere in lowlands, from dry to wet. Sometimes cultivated, and often seen on farms or along roads. Recognition: I. bella is similar but has somewhat smaller leaflets, less impressed veins, and its twigs are not square; see also I. chocoensis. I. spectabilis is more widely distributed than either of those species.

Fabaceae—Mimosoideae

Inga sapindoides

Inga spectabilis

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Fabaceae—Mimosoideae Inga thibaudiana

Inga thibaudiana

Inga thibaudiana. Leaflets in four to seven pairs,

Inga umbellifera

Inga umbellifera. Leaflets in two or three pairs, smooth and hairless. The rachis is winged. Fruit are short, with only two seeds. d istribution: Eastern half of Panama and throughout Costa Rica, in moist to wet zones, often at forest edge or streamsides. Recognition: I. marginata is similar, but always has two leaflet pairs, whereas I. umbellifera will usually have three pairs at least somewhere on the tree.

with soft hairs underneath creating a light green color. Veins on the underside are reddish. No wing on the rachis. Glands are like little cups, taller than in most other Inga. d istribution: Seen widely in wet lowlands and some lower montane sites throughout both countries. Sometimes along roads and streams. Recognition: Easy to confuse with I. cocleensis and I. multijuga, which are very similar but have smaller leaf glands.

Inga umbellifera

Fabaceae—Mimosoideae

Inga thibaudiana

Inga umbellifera

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Fabaceae—Mimosoideae Inga venusta

Inga venusta

Inga venusta. Leaflets in three pairs, with the rachis winged. There are soft red hairs on twigs, rachis, veins, and young leaves, but older leaves are smooth. d istribution: Mainly a lower montane species throughout, but also in wet lowlands. Recognition: Like I. sapindoides, but the latter has barely visible yellowish hairs and a very narrow wing.

Inga vera

cific coast in Panama and Costa Rica, with scattered records in wetter lowlands. A common tree in secondary forest, roadsides, and streamsides. Very common along roads on the Pacific half of the Canal Area. Recognition: Other Inga with so many leaflets do not have wings, but when I. vera has only three pairs, it looks like I. venusta or I. sapindoides.

Inga vera

Inga vera. Not illustrated. Leaflets in three to six pairs, or more, the outer pair much larger. Rachis is winged, and there is dense red pubescence on leaves, twigs, and the rachis, though mostly in younger leaves. Fruit is thick but constricted between seeds. d istribution: Largely in dry areas of the Pa-

Unfortunately, there are many more Inga in Panama. Quite a few are very sparsely known, and we believe we have covered those most often seen. Realistically, however, you will have to consider that a species you find may not be among those we describe here, given the total of 55 species in the country. On the other hand, everyone quickly learns the genus from the rachis glands.

Mimosa tenuiflora

Mimosa tenuiflora

Mimosa tenuiflora (herrero). A small tree of dry

savannas. The trunk has low branches, and the bark

is red. Branches have small, sharp spines. Leaves are bipinnate and pinnules are tiny. d istribution: Only known in the driest zone of Panama, in pastures and open areas. Recognition: A typical species of arid zones, with spines and tiny leaflets. Prosopis (Mimosoideae) is quite similar, but its spines are much larger. See Caesalpinia and Parkinsonia of the Caesalpinioideae, but they lack spines. A second species of Mimosa, M. pellita, is a shrub or treelet with similar small leaflets, occurring in swampy areas of the Pacific coast in both countries.

Fabaceae—Mimosoideae

Inga venusta

Mimosa tenuiflora

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Fabaceae—Mimosoideae Parkia nitida

Parkia nitida

Parkia nitida (dormilón, pellejo de sapo). A tall wet-forest tree. Trunk is light brown, smooth but with

small white lenticels. Leaves are bipinnate, but unusual for Fabaceae, the leaves are opposite (or very close to opposite, a condition called subopposite). Pinnules are tiny. There is one small gland per leaf, on the petiole just below the basal pair of leaflets. Flowers are white, typical mimosoid balls of protruding stamens. d istribution: A rare Caribbean slope species in Panama; not known in Costa Rica. Recognition: Leaves are very like those of the two species of Enterolobium, but their habitats are completely different. With leaves in hand, Parkia has its leaf gland in a different place, and its leaves are opposite.

Pithecellobium unguis−cati

Pithecellobium unguis-cati

Pithecellobium unguis-cati (espino carbón). A medium-sized tree of dry areas. It often has low branches and a wide crown. Branches have numer-

ous knobs, some of which have sharp spines. Leaves are bipinnate, with two pairs of two pinnules; there is a circular gland on the rachis between the leaflets. Flowers are white, on a spike. Pods are twisted and constricted between seeds. d istribution: Only found in the driest zones of Panama and Costa Rica, generally close to the Pacific coast. Recognition: Two pairs of two pinnules is an unusual character; see Zygia or Hymenaea (Caesalpinioideae). Two other Pithecellobium species are found in similar habitat. Both have two pairs of two pinnules and spines.

Fabaceae—Mimosoideae

Parkia nitida

Pithecellobium unguis-cati

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Fabaceae—Mimosoideae Prosopis juliflora

Prosopis juliflora

Prosopis juliflora (algarrobillo, aromo, manca caballo, mesquite). A small tree of the dry zone. Branches have large, sharp spines in pairs. Leaves are bipinnate with small, narrow pinnules. Flowers are yellow, in spikes. d istribution: Only in the driest zone of Panama and Costa Rica. Fairly common at Sarigua National Park. Recognition: Several small trees of dry regions have similar leaves; this one has the largest spines (spines of Mimosa are much smaller). This is related to the well-known mesquite trees of southwestern deserts in the United States.

Pseudosamanea guachapele

Pseudosamanea guachapele

Pseudosamanea guachapele (guachapalí,

guábilo, frijolillo). A tall tree of regrowth. Bark is lightcolored, deeply fissured into vertical plates. Leaves are bipinnate, pinnules moderate in size, round at the

tip; outer pinnules are much larger than the basal pairs. There is one gland per leaf, on the rachis between the outermost leaflets. Flowers are white pompoms. d istribution: Mostly in the agricultural zone of Pacific Panama and nw. Costa Rica. Very common in secondary forests, forest edge, and roadsides in the vicinity of Panama City. Recognition: Very similar to Samanea saman. Both have bipinnate leaves of about the same size, and both occur in open areas. But Samanea has purple flowers, often present, and is frequently a pasture tree, with huge branches near the ground. The other bipinnate mimosoids have tiny leaflets.

Fabaceae—Mimosoideae

Prosopis juliflora

Pseudosamanea guachapele

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Fabaceae—Mimosoideae Samanea saman

Samanea saman

Samanea saman (guachapalí, cenízaro, rain tree).

A large pasture tree. The trunk is huge, and the bark light colored and deeply fissured. Crown is very wide, easily wider than the tree is tall. Leaves are bi-

pinnate with moderate-sized pinnules all of similar size. Each pair of leaflets has a rachis gland between. Flowers are purple pom-poms. Pods are like those of Pseudosamanea guachapele. d istribution: In pastures of the dry Pacific coast, never in forest. Widely planted around the world as an ornamental. Recognition: Very similar to Pseudosamanea guachapele in both leaves and bark. But Pseudosamanea does not fork so close to the ground or have such a broad crown, and is mainly a tree of secondary forest and edge, not pasture (though it can grow in open areas). Samanea typically has its purple flowers throughout the wet season. Enterolobium cyclocarpum is a big pasture tree in the same habitat, but has much different leaves and bark.

Zygia latifolia

Zygia latifolia

Zygia latifolia (guabito de río, guabito cansa boca). A small swamp tree. Leaves are bipinnate, but with an unusual form that requires some study. Each leaf forks into two main leaflets, close to the branch.

Each of the two leaflets has five pinnules, two pairs plus an extra one very close to the base. At first glance they probably will not look bipinnate. Veins of the pinnules are asymmetric. There is a gland on the leaf rachis between the two main leaflets. The flowers are mostly pink stamens, and the pods are long, flattened, curved, and eventually split open to reveal seeds. d istribution: Only in swampy areas along the Caribbean coast. At the mouth of Chagres River it is fairly common. Recognition: At a glance, the leaves look like those of Inga laurina, but look closely at the layout of the leaflets, which only Zygia has. See Z. longifolia, but it has fewer pinnules.

Fabaceae—Mimosoideae

Samanea saman

Zygia latifolia

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Fabaceae—papilionoideae Zygia longifolia

Zygia longifolia

Zygia longifolia (guabito de río, guabito cansa boca, pichindé). A small to medium-sized tree of streambanks. The trunk base can be thickened, with many roots spreading outward. Bipinnate leaves are similar to those of Z. latifolia, but have only three pinnules per leaflet. There are glands on the leaf rachis between the leaflets and between the pinnules. Flowers and fruits as in Z. latifolia. d istribution:

Throughout Panama and Costa Rica, always near streams, sometimes on bare rocks with roots touching the water. Recognition: Best learned by its habitat. Leaves might be confused with those of Inga laurina or Z. latifolia, but upon close inspection are easily recognized. Panama has three more Zygia species, all of wet forests. Some have the odd arrangement of Z. latifolia, others not. They all have rachis glands like those of Inga, but are bipinnate. Four other genera of Mimosoideae are found in Panama, each with a single species: Balizia, Chloroleucon, Piptadenia, and Zapoteca. C. mangense is another tree of the dry Pacific zone with tiny leaflets like those of Acacia. The other trees are rare, especially so in Panama.

Acosmium panamense

Acosmium panamense

papilionoideae (subfamily within Fabaceae). This is the biggest and most widespread

of the three Fabaceae subfamilies, and the one including the most familiar plants: beans, peas, clover, and alfalfa. With 11,500 species in the world, it would by itself rank as the fourth biggest plant family, slipping behind the Rubiaceae (which the Fabaceae as a whole surpasses). There are about 4000 species in the American tropics. Many are trees, but larger numbers are lianas or herbs. There are 66 native tree species in Panama checklist, fewer than Mimosoideae but more than Caesalpinioideae. We illustrate 23 and describe 4 more.

Like the other groups of Fabaceae, Papilionoideae (nearly) always have compound leaves, and the base of the petiole is swollen and cylindrical. Unlike mimosoids, none are bipinnate. The flowers are most characteristic: the pea flower, with petals arranged into a beaklike structure (called the keel) and the reproductive parts hidden inside; two other petals behind are called wings. Most species have legumelike pea pods, but many trees have winged seeds and there are a few other fruit forms.

Acosmium panamense (frijolillo, malvecino). A

medium-sized tree of drier areas. Leaves are odd-pinnate, generally with 9–11 leaflets that are alternate to almost opposite along the rachis; the tip of each leaflet has a tiny notch. The upper side is a dark shiny green, and the underside lighter green. Flowers are white, seed pods winged. d istribution: Along the Pacific coast around Panama City and in Costa Rica. Recognition: See Dalbergia retusa, which also has notched leaflets.

Fabaceae—papilionoideae

Zygia longifolia

Acosmium panamense

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Fabaceae—papilionoideae Andira inermis

Andira inermis

Andira inermis (almendro de río, harino, quira). A tall forest tree. The trunk is straight and cylindrical, with small buttresses. Foliage is dense. Leaves are bright, shiny green, even-pinnate, with nine or more

leaflets that are opposite or nearly opposite one another (subopposite). At the base of each leaflet is a single, tiny, pointed stalk called a stipel. Flowers are purple; fruits are fleshy, not pods. d istribution: Very widespread, in all habitats throughout, often along rivers. Occurs in mature forest and in open savanna areas. Recognition: The odd little stipels are the key. Similar shiny leaflets are also found in Acosmium, Dalbergia, Platymiscium, and Vatairea. Lennea also has stipels, but its leaflets are rounded and not shiny. A second species of Andira with very similar leaves and larger fruits is known only from e. Panama.

Clitoria glaberrima

Clitoria glaberrima

Clitoria glaberrima. A small tree of dry regions.

The leaves have three pointed leaflets, dark green above, light bluish below. The outer leaflet is larger than the other two. Flowers are purple. d istribution: Along the Pacific coast of c. Panama and Costa Rica; also on limestone sites near the Canal (Madden Dam). Recognition: See Erythrina, which also has three leaflets.

Fabaceae—papilionoideae

Andira inermis

Clitoria glaberrima

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Fabaceae—papilionoideae Dalbergia retusa

Dalbergia retusa

Dalbergia retusa (cocobolo). A medium-sized tree of drier areas. The trunk is sometimes straight and tall, but more often irregular and branched near the base. Bark has vertical fissures. Leaves are oddpinnate, usually with 11–13 leaflets that are opposite or subopposite. They are light green or whitish on the underside, rounded, and the tip has a small notch. Flowers are white. Fruits are flat, winged pods with one to three seeds whose outlines are clear. d istribution: Found only on the drier half of the

isthmus. Common where dry forests remain in the driest zone of Panama and Costa Rica. Recognition: Acosmium is very similar, also with the small notch in each leaflet, but leaflets of Acosmium are not so white on the underside and are usually closer to alternate along the rachis. Andira is also similar, but lacks the notch and has opposite leaflets. There are many other papilionoids that have somewhat similar leaves: Diphysa, Diplotropis, Platypodium, and Vatairea. Cocobolo is well known for its carvings in tourist shops in Panama. Six more tree species of Dalbergia are known in Panama. Two are Caribbean coast species, and three others are very rare. The sixth, D. brownei, is often a liana but can grow on its own. One species, D. monetaria, has simple leaves, unusual among the Fabaceae. Others have compound leaves resembling those of D. retusa.

Diphysa americana

Diphysa americana

Diphysa americana (macano, cacique). A treelet

of dry and open zones. Leaves have small, rounded leaflets that alternate along the rachis, with undersides light bluish in color. Flowers are bright yellow. d istribution: Very common in open areas of the Pacific slope, often along roads; easy to see along the Pan-American Highway when flowering. Not generally inside the forest. Recognition: Platypodium has similar leaflets, but is a big forest tree. Check also Cassia moschata (Caesalpinioideae).

Fabaceae—papilionoideae

Dalbergia retusa

Diphysa americana

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Fabaceae—papilionoideae Diplotropis purpurea

Diplotropis purpurea

Diplotropis purpurea. A tall tree from wet for-

ests. The trunk is large and straight, with modestsized buttresses in big trees. Leaves are odd-pinnate, with medium-sized, pointed, rather narrow leaflets alternating along the rachis. Flowers are light pink, in dense spikes. d istribution: We discovered it in Panama in 2003 in plots on Santa Rita east of the Canal. Likely to turn up elsewhere on the Caribbean slope, including Costa Rica. Recognition: The pointed leaflets are rather like those of Dipteryx, but Dipteryx has narrow green wings on the rachis.

Dipteryx oleifera

Dipteryx oleifera

Dipteryx oleifera (almendro, almendro de mon-

taña). A very tall forest tree. The tallest tree at Barro Colorado I.—53 m—is this species. The trunk is immense, straight, swollen at the base into rounded buttresses. The bark is yellowish and granular. Leaves usually have 11–13 leaflets that alternate along the rachis; often, the rachis extends beyond the last leaflet. The leaflets are asymmetric, with the leading half wider than the trailing half. The petiole and leaf

rachis are winged, although the wing is narrow. Flowers are purple, produced above the foliage in June (at Barro Colorado I.), conspicuous at a distance. Fruits have a single, hard seed with a thin layer of brown pulp surrounding it; they are typically found on the ground with the pulp gone, and often with a hole chewed in the wall. During December and January, seeds are a staple for mammals at Barro Colorado I., and agoutis can frequently be heard gnawing on the seeds at the base of big trees. d istribution: A conspicuous canopy emergent at Barro Colorado I. and Soberania, where it is fairly common. Otherwise widespread but sparse along the Caribbean half of the isthmus. Recognition: An oft-studied tree at Barro Colorado I., it is easy to learn either from the trunk and yellowish bark, or the very distinctive leaves. Beneath big trees, the leaf rachis can usually be located, recognizable from the small wing even if the leaflets are gone.

Fabaceae—papilionoideae

Diplotropis purpurea

Dipteryx oleifera

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Fabaceae—papilionoideae Erythrina costaricensis

Erythrina costaricensis

Erythrina costaricensis (pito, machetito, palo

santo). A small forest tree with brilliant red flowers.

The trunk is short, usually leaning or bent, and has sharp spines. Leaves have three heart-shaped leaflets, the outer somewhat larger, light green or bluish beneath. Flowers are long, narrow tubes that barely open; they appear when the tree is leafless. d istribution: Widespread, but best known in the Canal Area, where it often grows immediately adjacent to small forest streams; common along the Pipeline Rd. streams. Flowers stand out in the dark forest understory. Recognition: Easy to learn. Trifoliate leaves are rare, and this is obviously Fabaceae (swollen petiole base) and has spines. See Clitoria.

Erythrina fusca

Erythrina fusca

Erythrina fusca (palo de bobo, palo santo, pito,

gallito). A medium-sized tree of lakeshores. The trunk can be large, but is usually branched near the ground; it has big, curving buttresses. There are large flat spines on the bark, and branchlets have small, sharp spines. Leaves have three leaflets, with small, rounded glands on the petiole near the base

of each. Flowers are large, orange, with a clawlike shape. They are produced conspicuously in December and January, and many birds are attracted to the nectar. The fruit is a long, twisted pod. d istribution: A tree of the margins of lakes, large rivers, or swamps; large stands of this species dominant many swamps in Panama. The shores of the Canal and Gatun Lake are lined with it. Away from water, it also grows in farmland, sometimes in fencerows. Recognition: Should be unmistakable, with or without flowers. Eight more Erythrina species are known in Panama, all with similar flowers and leaves. E. gibbosa is a montane and wet-forest species; E.rubrinervia is in the dry zone. Others are rare.

Fabaceae—papilionoideae

Erythrina costaricensis

Erythrina fusca

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Fabaceae—papilionoideae Gliricidia sepium

Gliricidia sepium

Gliricidia sepium (balo, mata ratón, madero ne-

gro). A medium-sized farm and fence tree. The trunk

is much branched, and the crown wide and dense. Branches are long, densely packed with long, alternate, odd-pinnate leaves carrying many pointed, opposite leaflets. Flowers are purple to pink, in dense clusters that are frequently present. d istribution: Generally around human habitation, especially on the Pacific slope, and often used in fencerows. Not in the forest. Perhaps spread widely by people, so its natural range obscure. Recognition: Dense stands along fences and roads are learned from a distance. Other papilionoids with similar leaflets do not have so many leaves and do not grow in fences.

Lennea viridiflora

Lennea viridiflora

Lennea viridiflora (algarrobillo). Not illustrated. A medium-sized tree of dry zones. Leaves are odd-pin-

nate, with seven to nine rounded, light green leaflets that are opposite along the rachis; each leaflet has a tiny notch at the tip. There is a tiny projection (stipel) at the base of each leaflet. Flowers are yellow and fruit a typical legume pod. d istribution: Fairly common in pasture and scrub of the Pacific coast in c. Panama; also in n. Costa Rica and on limestone around Madden Dam. Recognition: See Acosmium and Dalbergia, which also have notched leaflets and occur in the dry zone. Andira inermis has similar stipels but darker, more numerous, pointed leaflets.

Lonchocarpus heptaphyllus

Lonchocarpus heptaphyllus

Lonchocarpus heptaphyllus (chaperno, zorro).

A medium-sized tree of moist to wet zones. The trunk is straight, and bark black. Leaves are odd-pinnate, typically with seven to nine pointed leaflets in oppo-

site pairs; the terminal leaflet is enlarged. The leaf underside is light blue-green, and veins are yellowish. Flowers are pink; fruits are flattened pods. d istribution: Lowlands in wet to moist or even lower montane forest; quite common at Barro Colorado I. and Soberania. Recognition: Lonchocarpus is most like Ormosia in general form; both are odd-pinnate with strictly opposite leaflets. Platymiscium has similar leaflets but opposite leaves. Andira also has pointed, opposite leaflets, but they lack the bluish leaf undersides and are more numerous. Other papilionoids have alternating leaflets. The various species of Lonchocarpus differ in detail.

Fabaceae—papilionoideae

Gliricidia sepium

Lonchocarpus heptaphyllus

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Fabaceae—papilionoideae Lonchocarpus minimiflorus

Lonchocarpus minimiflorus

Lonchocarpus minimiflorus (frijolillo, malveci-

no). Similar to L. heptaphyllus, but with leaflets more numerous and smaller, and veins not so yellow. Flowers are dark purple; fruits like those of L. heptaphyllus. d istribution: Dry-forest areas of the Pacific slope; also common on limestone near the Madden Dam in the Canal Area. Recognition: See other Lonchocarpus and Ormosia, all of which have opposite leaflets.

Lonchocarpus sericeus

Lonchocarpus sericeus

Lonchocarpus sericeus (chaperno, guabito).

Similar to other Lonchocarpus, but leaflets are small, round or blunt-tipped, and have soft reddish hairs on the underside. d istribution: Largely in the dry zone of the Pacific coast, but not seen often; we know it along streams and swampy areas in Herrera. Recognition: See other Lonchocarpus and Ormosia, all of which have opposite leaflets.

Lonchocarpus velutinus

Lonchocarpus velutinus

Lonchocarpus velutinus (chaperno). Not illustrated. Similar to other Lonchocarpus, but with fine reddish hairs on twigs. Leaflets are fairly small and have a dense coat of pubescence beneath.

d istribution: Largely in the dry, agricultural zone of c. Panama, and can often be seen along roads. Recognition: See other Lonchocarpus and Ormosia, but the fuzzy leaves set this apart from most. The rare O. amazonica is quite similar but has a nonoverlapping range. Panama has 15 species of Lonchocarpus. L. atropurpureus is known in wet forests of the Canal Area. L. oliganthus is a widespread lower montane and wetforest species. Most of the remaining are scarcely seen. The genus can usually be learned from the opposite leaflets and tendency toward lightish leaf undersides.

Fabaceae—papilionoideae

Lonchocarpus minimiflorus

Lonchocarpus sericeus

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Fabaceae—Papilionoideae Myroxylon balsamum

Myroxylon balsamum

Myroxylon balsamum (bálsamo, bálsamo de tolú, sándalo). A tall tree of wetter zones. Big trees

warty. Twigs are covered in small white lenticels and have ribs running their lengths. Leaves are odd-pinnate and generally have 8–10 pointed leaflets alternate (or subopposite) along the rachis. When held to light, the leaflets have small translucent points or lines (use a hand lens). Flowers are white; fruits winged and large. Distribution: Sparsely in wet and lower montane areas, but rare outside the Darien. Recognition: See Acosmium and Dalbergia, with similar, alternating leaflets, but with notched tips. Vatairea also has similar leaves, but a much different trunk.

have small buttresses; the bark is light brown and Ormosia amazonica

Ormosia amazonica

Ormosia amazonica (frijolito de la suerte, cora-

lillo, peronil). A medium-sized wet-forest tree. Branchlets have dense, fine, reddish pubescence. Leaves are

smooth, dark green above, light below with reddish veins from a dense layer of hairs, odd-pinnate with five to nine rounded leaflets in opposite pairs. Leaflet borders are rolled downward slightly. Flowers are purple, and fruits a short pod densely covered in red hairs. The seeds within are half red, half black. Distribution: Known largely from our own inventories, and only at a few lower montane and wet-forest sites. Recognition: See Lonchocarpus, Ormosia, and Platymiscium. Much like Lonchocarpus velutinus, which also has red hairs and similar leaves. Compare to L. heptaphyllus as well.

Fabaceae—Papilionoideae

Myroxylon balsamum

Ormosia amazonica

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Fabaceae—Papilionoideae Ormosia coccinea

Ormosia coccinea

Ormosia coccinea (alcornoque, frijolito de la suerte, cabresto, coralillo, peronil). A large forest

tree. Trunk is straight, cylindrical, with small buttresses when large. Bark is light brown, and there are horizontal ridges ringing the trunk. Leaves are odd-

pinnate with five to nine leaflets that are strictly opposite. Top side of leaf is dark, smooth green, but the underside is pubescent with reddish veins. The leaf rachis and the petiole are also pubescent. Flowers are purple. Fruit is a small pod, flattened, pointed at one end; within are one or two seeds. The seeds are black on one half, bright red on the other. Distribution: Widespread from moist to wet and lower montane forest, on both slopes. Recognition: See Andira, Lonchocarpus, Ormosia, and Platymiscium. Of those, the most likely to be confused are L. heptaphyllus, which has yellowish veins, and O. amazonica and L. velutinus, which have pubescence. The other common Ormosia, O. macrocalyx, has no pubescence. The striking seeds are used for jewelry.

Ormosia macrocalyx

Ormosia macrocalyx

Ormosia macrocalyx (alcornoque, coralillo, peronil, cabresto, palo de collar, janeiro, nené, conejito colorado). A tall tree with a straight, cylindrical trunk, buttressed at the base when large. Leaves are odd-pinnate, with fairly large, dark shiny green leaflets lacking pubescence and opposite one another. When very small, saplings can have opposite leaves. Flowers pink to purple. Fruits are small pods in which there are bright, entirely red seeds. Distribution: Mostly known in our own inventories

in c. and w. Panama, especially in the Canal Area, where found from Pacific to Caribbean; few records from Costa Rica. Generally a tree of mature forest, but can be seen along roads and in towns and sometimes planted as an ornamental; there are a number of large individuals in Gamboa. Recognition: The compound leaves with large, shiny leaflets are pretty distinctive. A few bright red seeds can often be found beneath big trees, and these cinch the identification. Lonchocarpus heptaphyllus and O. coccinea have leaflets of similar size and shape, but they are pubescent and not shiny. Juveniles of Platymiscium pinnatum can be confused with O. macrocalyx; both can have opposite leaves. Three more species of Ormosia are known in Panama, all rare and one restricted to the Darien. There may be identification problems among several of the species. All have odd-compound leaves with opposite leaflets.

Fabaceae—Papilionoideae

Ormosia coccinea

Ormosia macrocalyx

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Fabaceae—Papilionoideae Platymiscium pinnatum

Platymiscium pinnatum

Platymiscium pinnatum (quira). A tall forest tree. Trunk is straight, cylindrical; large individuals have small rounded buttresses at the base. The bark is light brown to gray, and in larger trees has vertical fissures and thick plates. Leaves are opposite, which is very rare for the Fabaceae. They are odd-pinnate, and each has five to seven fairly large opposite leaflets. That is, leaflets within a leaf are opposite and

the entire leaves are opposite too. Flowers are yellow; fruit is a large, winged seed. Distribution: A widespread tree from dry to moist and even wet areas. Fairly common in remnant dry forests and sometimes open savannas of the Pacific coast. Recognition: Opposite, compound leaves are very rare. Only in the Bignoniaceae are they typical, but nothing in that family has leaves resembling those of Platymiscium. Turpinia occidentalis (Staphyleaceae) is likewise opposite-compound, but its leaflets are toothed. Lonchocarpus and Ormosia coccinea have leaves like those of Platymiscium, but they are alternate. As a caution, though, juvenile O. macrocalyx can have opposite leaves, but its leaflets are shiny. The quira has heavy, hard wood, prized for furniture and tools. A second species of Platymiscium is known only in e. Panama. Its leaves are very similar.

Platypodium elegans

Platypodium elegans

Platypodium elegans (carcuera, costilla, arbol soga, canalua, canaleto). A large forest tree. The trunk is bizarrely fenestrated. In big individuals, the fenestrations penetrate almost to the center of the trunk, and there are usually gaps you can look through to the other side. Bark is dark brown. Leaves are alternate, compound, with small, rounded leaflets, up to 20 or more per leaf. The leaflets are not quite opposite one another, and have a notch at their tips. Flowers are yellow. The fruit is a large wing, with a seed at one end; it

starts green but turns brownish. Distribution: Uncommon but widespread through Barro Colorado I. old-growth forest and at Soberania. Otherwise known sparsely in Panama, not in Costa Rica. Usually only giant trees are encountered; saplings are scarce and mostly confined to large forest gaps. Recognition: As a large tree, the bizarre trunk is frequently taken for a strangler fig; note that sometimes after a host tree dies, a strangler stands on its own. But most stranglers have a host tree underneath, and regardless, stranglers have light-colored trunks that are rounded, quite unlike Platypodium. Check Minquartia (Olacaceae) and Aspidosperma (Apocynaceae), also with fluted trunks, but no other nonfig has such deep invaginations as Platypodium. Even in tall forest, the small leaflets of Platypodium are easy to see; they resemble those of Cassia moschata (Caesalpinioideae) and another big papilionoid, Myrospermum frutescens (not illustrated).

Fabaceae—Papilionoideae

Platymiscium pinnatum

Platypodium elegans

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Fabaceae—Papilionoideae Pterocarpus officinalis

Pterocarpus officinalis

Pterocarpus officinalis (sangre de gallo, cricamola, suela). A tall swamp tree. Big trees have

enormous buttresses, extending well up the trunk. The bark is light colored, smooth. Leaves are shiny green above and below, odd-pinnate, with seven to nine leaflets alternating along the rachis. Flowers are yellow, pods circular and flattened. Distribution: Common and sometimes the dominant tree in swamps along the Caribbean coast and Pacific coast rivers of the Darien and the Osa Peninsula. Occasionally grows upland, but usually not far from the coast. Recognition: The three Pterocarpus are all similar; this one is best known by habitat.

Pterocarpus rohrii

Pterocarpus rohrii

Pterocarpus rohrii (sangre de gallo, cricamola, suela). A tall forest tree. Trunk straight, cylindrical. Large individuals have high and thin buttresses. The bark is smooth and yellow with white splotches. Leaves are odd-compound, with five to nine leaflets that alternate along the rachis. Leaflets are shiny green. Flowers are yellow. The fruit is a large, circular, winged seed, greenish but turning light brown. Distribution: Wet to moist and lower montane for-

ests. Quite common near the Canal and in parts of Costa Rica, but sparse elsewhere. Very common in the old forest at Barro Colorado I. Recognition: Alternating leaflets of Pterocarpus resemble those of Acosmium and Dalbergia, but those two have blunttipped leaflets. Other legumes that are big trees have opposite leaflets. The unrelated Picramnia latifolia of the Picramniaceae, a family we do not cover at all (see Appendix 3), is remarkably similar to Pterocarpus rohrii, including a very legumelike cylindrical petiole base. Picramnia is no more than a few meters tall, but that is no help separating juveniles. You can chew the leaflets; those of Picramnia are very bitter. Pterocarpus has one other species in Panama, P. acapulcensis. It has small, opposite leaflets, bluish underneath, and is known from only a few sites on the Pacific coast of Panama.

Swartzia panamensis

Swartzia panamensis

Swartzia panamensis (cutarro, malvecino). Not illustrated. A tall wet-forest tree. Large trees have

small buttresses. Bark is light, yellowish. Leaves are odd-pinnate, with five to seven narrow leaflets that have greatly elongated tips. Flowers yellow, in hanging clusters. Fruits are flat, rectangular pods, smooth on both sides. Distribution: Caribbean wet forests in the Canal Area; also in Darien and the Osa Peninsula. Known in swampy areas at Soberania. Recognition: With just five opposite leaflets, reminiscent of Lonchocarpus. Leaflets are like those of Pterocarpus, but Pterocarpus has alternate leaflets. The exaggerated leaflet tip of S. panamensis is very distinctive.

Fabaceae—Papilionoideae

Pterocarpus officinalis

Pterocarpus rohrii

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Fabaceae—Papilionoideae Swartzia simplex

Swartzia simplex

Swartzia simplex (naranjita, naranjito, naranjo de

monte, limoncillo). A medium-sized forest tree. The trunk is often leaning, twisted, and branched low. Leaves are typically odd-compound, having three to five alternating leaflets. But there are also simpleleaved forms of this species, and sometimes the same tree will have both simple and compound leaves. Whether simple or compound, leaves are similar: shiny, smooth green, with a long, pointed tip,

and a leaf rachis (or petiole below the single leaf) with a very narrow green wing. Flowers are large and yellow. Fruits are bright orange and open to reveal a shiny black seed. Distribution: Very widespread; found in essentially all forest habitats from dry and wet to lower montane. Usually inside mature forest. Recognition: Does not immediately look like Fabaceae, because leaflets are few or the leaves are simple. But it does have the swollen petiole base typical of the Fabaceae. The faintly winged rachis is a good clue, easy to learn because the tree is encountered so often. The wing is reminiscent of leaves of Citrus (Rutaceae), hence the common names. Swartzia includes one other species in the region, but it is rare in w. Panama and s. Costa Rica. We recently discovered another species in the genus that we still have not identified.

Vatairea erythrocarpa

Vatairea erythrocarpa

Vatairea erythrocarpa (amargo-amargo). A tall wet-forest tree. Bark is brown to reddish, and big trees have spreading buttresses. Leaves are odd-pinnate, with 11–15 pointed, smooth leaflets that alternate (sometimes almost opposite) along the rachis. Flowers are light purple, and fruit is a large winged seed. Distribution: Known in the wetter forests around the Canal and nearby mountains; very rare in Costa Rica. Quite common near Pipeline Rd. and near Gamboa. Recognition: The numerous leaflets are quite similar to those of Andira inermis. Leaflets of Vatairea usually look alternate enough to avoid confusion with Lonchocarpus and Ormosia. Check also Acosmium, Dalbergia, and Myroxylon, leaflets of which are alternate or nearly so, and Pterocarpus,

which has fewer leaflets that are more obviously alternate. The trunk of big Vatairea can be confused with that of Tachigali versicolor (Caesalpinioideae). There are still another 14 genera of papilionoids we have not covered: Ateleia, Centrolobium (two), Crotalaria, Desmodium, Dussia (five), Fissicalyx, Hymenolobium, Lecointea, Machaerium, Muellera, Myrospermum, Paramachaerium, Piscidia, and Sesbania. They have a single species each except the two indicated with parentheses. The genus Dussia is important in the Neotropics, and has a couple fairly common wetforest species similar to Lonchocarpus and Ormosia in leaf form. A few of the rest can also be common: Fissicalyx is often seen in forests around Panama City, and has 11 narrow, pointed leaflets and yellow flowers; Machaerium is a widespread, spiny dry-forest tree with many rounded leaflets; Hymenolobium is a tall tree of wet forests in c. Panama and n. Costa Rica with leaves that have more than 30 rounded leaflets; and Desmodium cajanifolium is a weedy treelet found everywhere in South America and common in open areas of Panama. The remaining genera are seldom seen.

Fabaceae—Papilionoideae

Swartzia simplex

Vatairea erythrocarpa

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Fagaceae

Fagaceae

Quercus insignis

Quercus insignis

This is the oak family, with around 800 tree and occasionally shrub species. It is the dominant family of many forests and savannas throughout north temperate regions. In tropical America, there are about 150 species, most in mountains of s. Mexico at the margin of the tropics. In Panama and Costa Rica, oaks are almost exclusively in mountains above 1500 m. Nine species are known in Panama, and we cover one here.

Quercus insignis (roble, roble de montaña, enci-

peeling off in small pieces. Branchlets have fine reddish hairs. Leaves are toothed, simple and alternate, bunched at the end of the branchlets. The leaf underside is lightish blue but has prominent yellow veins and is covered with dense, fine hairs. The fruit is an unmistakable acorn. Distribution: Mostly in high mountains of far w. Panama and throughout Costa Rica. At that elevation, oaks are sometimes dominant. Recognition: No one will fail to recognize the acorns. Just like northern species, the leaves are bunched at the branch tips. Compare to Alnus (Betulaceae), which is also found at high elevation and has similar venation and leaf teeth. Eight more oak species are known in Panama, all from western highlands and all more widely distributed in Costa Rica, where there are more high mountains. One species is found in the Darien. Several have coarsely toothed leaves like those of Quercus insignis, and are not easy to separate.

no). Tall montane trees. Trunks are large, with bark

Hernandiaceae Gyrocarpus americanus

Gyrocarpus americanus

A small tropical family of trees, shrubs, and lianas. There are 60 species in the world, and 23 in tropical America. Panama has just four species, and we illustrate one and describe another. With such a small family with very different species, we omit the family description.

Gyrocarpus americanus (volador, helicóptero). Not illustrated. A rare species of the dry zone. Young leaves have three pointed lobes, but older leaves do not. In all leaves, there are three prominent veins arising from the base. Flowers are very small, white to yellowish, in tight but small clusters. Seeds have two long wings, thus the common name. Distribution: Found only in a few savanna locations in the driest parts of c. Panama and nw. Costa Rica; considered endangered in Panama. Recognition: The maplelike young leaves look like those of cotton (Gossypium). See also Heliocarpus americanus (Malvaceae—Grewioideae), which also has wide, lobed leaves.

Hernandia didymantha

Hernandia didymantha

Hernandia didymantha (zopilote). A mediumsized tree of wet forest. Leaves are shiny green, clustered toward the ends of the branches, round at the base but pointed at the tip, with three main veins arising from the base. There is no stipule. Crushed leaves have a fairly strong avocado-like odor (see the

family Lauraceae). Flowers are medium-sized, white to greenish, in terminal clusters. Fruits are fleshy, single-seeded, not winged. Distribution: Mostly in lower montane forests, but also in lowland wet forests of the Caribbean coast and the Osa Peninsula. Recognition: The wide, rounded leaves are very similar to those of Dendropanax arboreus (Araliaceae), but Dendropanax has petioles with widely variable lengths and also has a stipule. Two more Hernandia species are found in Panama. H. stenura is widespread in Costa Rica, less so in Panama, and more restricted to mountains than is H. didymantha; it also has rounded, clustered leaves. The other species is barely known.

Hernandiaceae

Quercus insignis

Hernandia didymantha

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Humiriaceae

Humiriaceae Humiriastrum diguense

Humiriastrum diguense

by the absence of traits is difficult: Humiriaceae lack odor, lack latex, do not have obviously clustered or regularly spaced leaves, and lack leaf punctations; when they also lack teeth and ridged stems, they are very likely to be confused.

Humiriastrum diguense (corocito, corozo). A

A small family of trees and shrubs, almost entirely from the American tropics. There are 65 species worldwide, with just 1 in Africa. Panama has four species, of which we cover three. Humiriaceae is not an easy family to learn. Leaves are simple and alternate, usually fairly small, and often but not always toothed. Their arrangement on branches is not especially striking, not densely clustered but also not regularly spaced. Twigs are sometimes ridged or square, but again not always. Neither flowers nor fruits are striking. Lowland trees with toothed leaves will most often be Salicaceae, but the latter usually have translucent leaf spots (punctations), which Humiriaceae lack. Any family known

tall wet-forest tree. Trunk is straight, cylindrical, buttressed at the base. Twigs are ridged. Leaves are dark green above and light below, smooth and hairless, simple, alternate, and finely toothed. In canopy trees, the leaves are small, but in juveniles they are larger and thinner. The petiole is very short and slightly swollen at the base. Flowers are small, greenish white, in terminal clusters. The fruit is single-seeded, fleshy, and avocado-shaped. Distribution: Known widely in wet and lower montane forests of c. Panama, plus the Osa Peninsula. Not especially common. Recognition: The combination of ridged branchlets with toothed leaves is distinctive. Look also for the short petiole that is swollen slightly at the base. The main lowland group with toothed leaves is Salicaceae, but Salicaceae usually have transparent dots in the leaves, and larger leaves with longer petioles.

Sacoglottis trichogyna

Sacoglottis trichogyna

Sacoglottis trichogyna (corocito, corozo). A tall

wet-forest tree. Trunk is straight, cylindrical, with small buttresses at the base. Twigs are ridged. Leaves

are small, simple, alternate, finely toothed; there are tiny black spots on the leaf border near the base. Small flowers are in mostly terminal clusters. Fruits are like those of Humiriastrum. Distribution: Found in wet forests of c. Panama and most of Costa Rica. Not especially common. Recognition: See the similar Humiriastrum diguense, which has shorter petioles. A second Sacoglottis species, S. ovicarpa, is known in wet forests of Panama but not Costa Rica. It is very similar, though with less-conspicuous teeth.

Humiriaceae

Humiriastrum diguense

Sacoglottis trichogyna

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Hypericaceae Vantanea depleta

Vantanea depleta

Vantanea depleta (chiricano, corocito). A tall wet-forest tree. Big trees have substantial buttresses. The bark is fissured so that pieces sometimes flake free. Twigs on large trees are not ridged, though they are on saplings. Leaves are simple, alternate, somewhat rounded, not toothed; the petiole is short. Flow-

ers are like those of Sacoglottis trichogyna. Distribution: Known mostly in c. Panama, in wet, moist, and lower montane forest. Common in forests of Pipeline Rd. and very common on the hill above Gamboa. Not known in Costa Rica. Recognition: Where common, the bark is recognizable. Leaves are similar to those of Sacoglottis, though without teeth; except in saplings, which have ridged twigs, Vantanea lacks an easy distinction. It resembles Maranthes panamensis (Chrysobalanaceae), but Maranthes has glands at the leaf base. Another species of Vantanea has been recorded in far w. Panama and the Osa Peninsula. It also has simple, untoothed leaves like those of V. depleta.

Hypericaceae Vismia baccifera

Vismia baccifera

A medium-sized family, with 500 species worldwide and about 175 in the New World. But most are herbs of temperate regions and tropical mountains. In Panama, just one genus includes small trees, and there are five species; we cover three. Hypericaceae is related to and sometimes included within the Clusiaceae, sharing opposite leaves and orange latex.

Vismia baccifera (pinta mozo, achiote, sangrillo, achiote tigre). A treelet of forest edge. The trunk is small and has low branches. Leaves are opposite, with a striking red underside composed of fine hairs. Broken leaves produce flowing orange latex. Flowers are star-shaped and white with purple lines, carried at the ends of branches. Fruits are spherical capsules that turn black and carry flower remnants. Distribution: Essentially all forest types throughout Panama and Costa Rica. Often on forest edge or roadsides in woods, and can be found along Pipeline Rd. in Soberania. Occasionally in openings in mature forest. Recognition: Check the other Vismia with red leaves, V. macrophylla. Other species with red leaf undersides (Luehea seemannii [Malvaceae—Grewioideae and Chrysophyllum cainito [Sapotaceae]) have alternate leaves.

Hypericaceae

Vantanea depleta

Vismia baccifera

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Hypericaceae

Vismia billbergiana

Vismia billbergiana

Vismia billbergiana (pinta mozo). A treelet often

of forest edge. Trunk is small, with low branches. Leaves are opposite, narrow and pointed, light green below. Broken leaves produce flowing orange latex.

Flowers are similar to those of V. baccifera, but are greenish and lack purple lines. Fruits are slightly smaller. Distribution: Same as that of V. baccifera: moist, wet, montane, often at forest edge. Recognition: Unlike the other two Vismia, lacks red under the leaves, so might be confused with many other species with opposite leaves. There are Apocynaceae with opposite leaves and latex, but they gush white latex. There are many other species with opposite leaves but no latex, including Rubiaceae and Myrtaceae; Vochysia ferruginea (Vochysiaceae) has leaves very similar to those of V. billbergiana and appears commonly on forest edge, but again, has no latex.

Vismia macrophylla

Vismia macrophylla

Vismia macrophylla (pinta mozo, achiote, san-

grillo). A treelet or small tree of montane forest edge.

Somewhat larger than the other Vismia, with a straight trunk so looking more treelike. Leaves are opposite, long, and conspicuously red beneath. Bro-

ken leaves produce orange latex. Flowers and fruits are like those of V. baccifera. Distribution: Abundant roadside tree in lower mountains, but also occurs in low-elevation moist and wet regions, in natural clearings within mature forest or on the edge. Fairly common along outer Pipeline Rd. Recognition: Large, opposite leaves with red underneath are unmistakable. Leaves of V. baccifera are much smaller. Two additional Vismia species are found in Panama. V. jefensis is endemic to Cerro Jefe and looks very similar to V. baccifera. The other is very common throughout the Canal Area, also along roadsides, with leaves like those of V. macrophylla but shorter.

Hypericaceae

Vismia billbergiana

Vismia macrophylla

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Icacinaceae

Icacinaceae Calatola costaricensis

Calatola costaricensis

A medium-sized and mostly tropical family, with 300 species of trees, shrubs, and lianas in the world. But there are only 50 American species, and Panama has just three species of trees. We cover one of these, and omit a family description.

Calatola costaricensis (jaguey). A medium-sized

cloud-forest tree. The trunk has low branches. Leaves are simple, alternate, sometimes but not usually

toothed. Secondary veins are prominent and widely spaced. There are tiny clusters of hairs in the vein axils on the leaf underside. Male flowers are tiny and white, on long spikes; female flowers are also small, white, on short stalks off small branches. Fruits are fairly large, fleshy, single-seeded. Distribution: Very widespread in lower montane and lowland wet forests throughout. Recognition: Not an easy species to learn. When teeth are present, it somewhat resembles species of Casearia (Salicaceae), and there are some of those in the mountains. Without teeth, it recalls Hieronyma oblonga (Phyllanthaceae), though Hieronyma has silvery scales on leaf undersides. Might also be confused with Lauraceae. Two other genera of Icacinaceae are known in Panama, Mappia and Poraqueiba, with one species each. Both are seldom seen and poorly known.

Lacistemataceae Lacistema aggregatum

Lacistema aggregatum

A small family, with just 15 species in two genera from tropical America. They are relatives of the Salicaceae, and like that family, leaves are alternate and often, though not always, toothed. We cover here two of the three species known in Panama.

Lacistema aggregatum (huesito). A very widespread treelet. Trunk is straight, cylindrical; branchlets have small swellings at the base of each leaf. Leaves are simple, alternate, regularly spaced along branchlets, in a flat plane. They can be untoothed or

slightly toothed. The stipules fall off readily, but they leave a clear scar, a ring that almost encircles the branchlet; that is, the scar wraps about three-quarters around, leaving an opening on the side opposite the leaf. Flowers are small, white, in small clusters of short spikes on the branches. Fruits are bright red berries. Distribution: A remarkably widespread treelet of forest understory, found in all forest zones except the highest mountains. Abundant in all forests of the Canal Area. Recognition: This is a difficult species to recognize at first, but it is common enough to encounter frequently and learn well, and the incomplete ring around branchlets is distinctive. When the leaves have teeth, it looks like a Salicaceae with its regularly spaced leaves in a flat plane, and it should suggest Casearia in that family. See Lozania pittieri, which is much less common. If teeth are not present and you don’t think of checking for the incomplete ring, Lacistema will be difficult to pin down.

Lacistemataceae

Calatola costaricensis

Lacistema aggregatum

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Lamiaceae

Lozania pittieri

Lozania pittieri

Lozania pittieri (huesito, garrotillo). A wet-forest treelet. The small trunk has low branches. Leaves are small, simple, alternate, regularly spaced and arranged in a flat plane. They have fine teeth, but this may be indistinct. Where secondary veins meet the

main vein on the leaf underside, there are tiny clusters of hairs; this requires a hand lens to see. Flowers are small, white, in long spikes from the branches. Distribution: Lower montane and wet forest throughout. Quite common along roads and trails at Cerro Campana. Recognition: Similar to Lacistema, but lacks the incomplete ring at the leaf bases and has tiny hairs on the leaf underside. Lacistema is more common in lowlands. See also Laetia thamnia or Casearia in the Salicaceae. A second Lozania species is very similar. It is known at lower montane sites in c. and w. Panama through Costa Rica.

Lamiaceae

Aegiphila anomala

Aegiphila anomala

One of the large plant families of the world, including 6500 species of trees, shrubs, herbs, and lianas, prevalent worldwide in temperate climates as well as the tropics. Tropical America has about 930 species. This is the mint family, with many species cultivated as spices; most of those are native to the Mediterranean, but sage of North American deserts is in the family. It is also the family of the Asian timber tree teak, which we describe below. Inside tropical forests, Lamiaceae is not an important family, having only a few species of trees and lianas. Panama has 21 species of trees in the family, plus teak. We cover five of these and illustrate four. All were formerly classified as Verbenaceae. Many know the mint family characters: square stems, opposite, toothed, aromatic leaves, and tubular flowers with two lips extended on the bottom. But these traits are not helpful in trees of the family, except for opposite leaves, which all species have. Most trees do not have teeth, though some do, and square

stems are likewise not reliable. Flowers of the trees are nearly regular and thus only vaguely mintlike. The tree Lamiaceae might be mistaken for Myrtaceae, but the latter should always have a clear collecting vein at the leaf margin. Several other minor opposite-leaf families (Celastraceae, Malpighiaceae, and Vochysiaceae) can be confused. Because the tree Lamiaceae are not common, you are likely to learn the individual species without paying attention to family-level traits.

Aegiphila anomala (tabaquillo). A medium-sized cloud-forest tree. Bark is gray. Twigs are square. Leaves are simple, opposite, narrow and pointed, without teeth; they tend to be clustered toward the ends of branches, with adjacent pairs perpendicular to one another. The leaf underside is much paler than above. The petiole is grooved on the upper side and has an expanded base. Flowers are white, with equal-sized petals. Fruits are single-seeded berries held in the calyx. Distribution: Mostly in lower montane forest from c. Panama through Costa Rica, but also found at lowland wet sites toward the Caribbean. Common in cloud forests at Cerro Campana. Recognition: Aegiphila in is not seen often and is not very distinctive. With clustered leaves in perpendicular pairs, this might be confused with Byrsonima (Malpighiaceae) but not Myrtaceae, which usually have all leaves in a flat plane. The pale leaf underside is a good character.

Lamiaceae

Lozania pittieri

Aegiphila anomala

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Lamiaceae

Aegiphila odontophylla

Aegiphila odontophylla

Aegiphila odontophylla (tabaquillo). A mediumsized montane tree. Bark is gray to brown. Twigs are square and have white lenticels. Leaves are simple, opposite, broad, and toothed, in perpendicular pairs and moderately clustered. Flowers and fruits are like those of A. anomala. Distribution: In mountains only, from w. Panama through Costa Rica. Recognition: Toothed leaves set in perpendicular pairs are mintlike, also resembling Acanthaceae. No other Lamiaceae (that are trees) have teeth.

Aegiphila panamensis

Aegiphila panamensis

Aegiphila panamensis (palomo, fruta de mono).

A small forest tree. Bark gray. Twigs are square or flattened. Leaves are simple, opposite, without teeth, pointed at both ends, spaced regularly and in a flat plane. The leaf underside is only slightly paler than above. Petiole is grooved above. Flowers are small, white, regular, in loose clusters held on long stalks all along the branch. Distribution: Found widely from dry to wet and lower montane forest, but rather sporadically. Seen regularly in forests of the Canal Area. Recognition: Leaves are held in a flat plane, unlike those of A. anomala. More likely confused with Celastraceae or Myrtaceae. Not an easy tree to recognize without experience.

There are 13 Aegiphila tree species in Panama, plus some lianas. The distinction can be blurred, though, because the trees of the genus can be liana-like. All have opposite leaves, mostly without teeth.

Tectona grandis (teak, teca). Not illustrated. The Asian timber tree. In Central America, it is a mediumsized tree, often with a forked trunk. Leaves are large to very large, simple, opposite, untoothed, in rotated pairs (one pair held perpendicular to the next), clustered toward branch tips. Flowers are small, white, regular, with no explanded petal, in large, loose clusters above the leaves. Fruits are dry, green capsules. Distribution: Planted widely near human habitation, in cities and rural areas, with many plantations in both Costa Rica and Panama. Recognition: Never a forest species, but it needs to be learned because it is so common in open areas and towns. The large leaves are reminiscent of those of the balsa (Ochroma [Malvaceae—Bombacoideae]) of the same habitat, but balsa leaves are lobed and alternate.

Lamiaceae

Aegiphila odontophylla

Aegiphila panamensis

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Lauraceae

Vitex cooperi

Vitex cooperi

Vitex cooperi (cuajado, flor azul). A tall tree of dri-

er areas. The trunk is straight, cylindrical. Bark is gray and sometimes peeling away in small pieces. Leaves are opposite, compound, with three to five untoothed leaflets; occasionally there is only one leaflet (and thus looking like a simple leaf). Flowers are purple, with one expanded, wrinkled petal. Fruits are berries that turn black. Distribution: Mostly in drier areas along the Pacific coast, but also seen

along the Caribbean coast. Recognition: The opposite, compound leaves are easy to spot and could only be confused with Tabebuia (Bignoniaceae), but the latter never has three leaflets. Four more Vitex species are known in Panama, all rare, and none are known in Costa Rica. V. cymosa is an immense tree of the Caribbean coast also having opposite, compound leaves and purple flowers. Three more genera of Lamiaceae are known among Panama’s trees: Callicarpa and Clerodendrum with one species each, and Cornutia with two. They are all small and weedy. Some species of Callicarpa and Clerodendrum are used as garden ornamentals. Another Asian tree, Gmelina arborea, has heartshaped opposite leaves and is sometimes planted in cities. It is sometimes called white teak.

Lauraceae

Beilschmiedia pendula

Beilschmiedia pendula

This is one of the major families of tropical trees. There are 2750 species in the world, nearly all trees, and about 1000 in tropical America. It is the family of avocado (Persea americana), a Central American species, and of cinnamon (Cinnamomum zeylanicum), an Old World species. Lauraceae is the fourth largest family of trees in Panama with 112 species; we illustrate 9 of those. Lauraceae have very consistent leaf characters, and this might suggest it is a good family to learn. Unfortunately, it is not. The problem is that the typical Lauraceae leaf pattern is rather nondescript and similar to that of many other families. The leaves are simple, usually alternate (but several species have opposite), untoothed, of moderate size, and an elliptical shape, all traits of the run-of-the-mill rainforest tree. But Lauraceae leaves have an avocado-like odor when crushed, akin to the odor of Annonaceae (the so-called ranalean odor), but unlike Myristicaceae, Lauraceae leaves are not regularly spaced or held in a flat plane. They tend to be clustered toward the end of branches and elevated above the horizontal. Moreover, branches of Lauraceae are neither precisely horizontal nor verticillate as in Annonaceae or Myristicaceae; instead, they tend to rise above the horizontal, and in saplings the upper branch ends above the top of the stem. Another good Lauraceae

trait is the pattern of secondary veins, which tend to be widely and often irregularly spaced, arching well forward. Finally, Lauraceae never have stipules. This all sounds good, but don’t get your hopes up, because the spicy odor is typically faint and sometimes not present at all, and the difference in leaf and vein arrangement among various families is subtle. Lauraceae fruits, however, are distinct, all being fleshy, one-seeded, and looking like miniature avocados. Flowers are always nondescript: white, tiny, and in loose clusters at the ends of branches. Lauraceae are common in all tropical forests, from lowland to montane, and you need to always have the family in mind.

Beilschmiedia pendula (aguacatillo, torpedo). A

tall old-growth tree. The trunk is straight, cylindrical, and slightly swollen into small buttresses at the base. The bark is light brown and peels off in patches to reveal large ovals of orange inner bark. Leaves are simple, alternate, bunched toward the end of branches. The leaf underside is a soft bluish color, and the color rubs away when scraped with a fingernail (the color comes from fine hairs that are indeed scraped away). Crushed leaves smell faintly like avocado. Fruits are like small avocados, with a large seed and a thin layer of pulp. Distribution: Found in moist, wet, and some lower montane forests through Panama and Costa Rica, though sporadic. Common throughout the Caribbean half of the Canal Area, and one of the dominant canopy species in old-growth forest at Barro Colorado I. Juveniles are common in the shaded understory, and big trees typically have dense concentrations of seedlings below the crown. Never an edge or a roadside species. Recognition: The rather large oval leaves with a bluish underside that can be scraped off are distinctive in juveniles. They resem-

Lauraceae

253

Vitex cooperi

Beilschmiedia pendula ble avocado leaves, but smaller. Nectandra lineata has similar leaves. In adult Beilschmiedia, the red and orange ovals on the trunk are very distinctive. Astronium graveolens (Anacardiaceae) also has light-colored ovals on its trunk, but not red or orange.

Panama has seven additional species of Beilschmiedia. They are rare in Panama, with many only in the western mountains; several are much more often seen in mountains and wet forests of Costa Rica.

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Lauraceae

Caryodaphnopsis burgeri

Caryodaphnopsis burgeri

Caryodaphnopsis burgeri (sigua blanco). A tall

wet-forest tree. The trunk is straight and cylindrical; bark is dark gray, sometimes with white spots. Leaves are opposite, dark green above but light green on the underside. Three main veins arise together at the leaf base and run parallel for the length of the leaf. The fruit is spherical. Distribution: Only known in wet forests of the Canal Area and s. Costa Rica. First dis-

covered in Panama at Pipeline Rd. in the mid-1990s, but rare there. Recognition: Opposite leaves with three main veins are unlike those of other Lauraceae in Panama. These characters should first make you think Melastomataceae, but melastomes have clearer tertiary veins and more often five parallel secondary veins. Caryodaphnopsis will be confused with the genus Strychnos (Loganiaceae, a family we do not cover), which has identical venation and opposite leaves. Most Strychnos are lianas, but there are rare treelets. We found a second species of Caryodaphnopsis, C. tomentosa, in Bocas del Toro but it has not been seen elsewhere in Panama or Costa Rica. It has similar leaves, but with a densely hairy underside.

Cinnamomum triplinerve

Cinnamomum triplinerve

Cinnamomum triplinerve (sigua, sigua blanca).

A tall tree of secondary forest. Big trees have small buttresses and sometimes sprouts near the base. Leaves are simple, alternate, with a faint spicy smell.

Two secondary veins arise from the main vein just above the leaf base, so there appear to be three main veins. Fruits are small. Distribution: Largely a species of edge and secondary forest, and seen widely in the driest areas through wet and lower montane regions. Common in the Canal Area. Recognition: Easy to learn these leaves, with their three prominent parallel veins. Six more Cinnamomum species are found in Panama. Most are montane and rare in Panama but more common in Costa Rica. Several of them do not have prominent secondary veins, and the similarity to C. triplinerve will not readily be recognized.

Lauraceae

Caryodaphnopsis burgeri

Cinnamomum triplinerve

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Lauraceae

Nectandra cuspidata

Nectandra cuspidata

Lauraceae that does not obviously fit the other genera we describe, always guess Ocotea but consider also Nectandra.

Nectandra cuspidata (sigua). A medium-sized

Genera Nectandra and Ocotea. These two groups dominate the Lauraceae of tropical America. Nectandra has 17 species in Panama, and Ocotea has 47, indeed, Ocotea is Panama’s fifth largest tree genus. Without flowers, they are very difficult or impossible to separate, and even the floral distinctions are subtle; experienced botanists have difficulty, and a book could be written about them. If you find a

tree of forest edge. The trunk can have small buttresses, and the bark has white spots. Leaves are simple, alternate, narrow, pointed at both ends. Secondary veins are very widely spaced. There are small clusters of hairs where secondary veins meet the main vein. Distribution: Pacific slope in c. to w. Panama, not Costa Rica; mostly at forest edge and roadsides. Very common in young forests from Panama City to Gamboa. Recognition: The narrow leaves and widely spaced secondary veins make the species reasonably distinctive, but like most Lauraceae, it does not stand out.

Nectandra lineata

Nectandra lineata

Nectandra lineata (sigua). A medium-sized tree

of forest edge. The trunk can have small buttresses. Twigs are ridged. Leaves are simple, alternate, narrow, slightly clustered toward the ends of branches and rotated relative to the branch and to each other, so not in a single plane (typical of Lauraceae). Veins

are yellow on the underside. There are tiny pits (called domatia) where secondary veins meet the main vein; these will require a hand lens to see. Tertiary veins are dense and parallel between the secondary veins. Flowers are white, in dense clusters above the leaves, larger than most Lauraceae flowers. Fruits are 1 cm and have a cup enclosing the base. Distribution: Widespread, mostly on the Pacific slope but also the Caribbean slope of the Canal Area; mostly at forest edge or in secondary forest. Common along wooded roads near Panama City, and the clusters of white flowers are easy to spot during December and January. Recognition: Prominent yellow veins and ladderlike parallel tertiaries are good marks in its secondary habitat.

Lauraceae

Nectandra cuspidata

Nectandra lineata

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Lauraceae

Nectandra purpurea

Nectandra purpurea

Nectandra purpurea (sigua). A medium-sized forest tree. Leaves are simple, alternate, smallish as Lauraceae go. They often have a long, pointed tip,

but not always. Secondary veins are few. There are small clusters of hairs where secondary veins meet the main vein. Distribution: Widespread, mostly in Caribbean wet forests to lower mountains. Common in moist to wet forests of the Canal Area. Recognition: Easy to overlook, because there is nothing striking about leaves or trunk, and without fruit, nothing clearly says Lauraceae. See Ocotea whitei, and among other families, check Phyllanthaceae and Euphorbiaceae. Several more fairly common Nectandra species are found in Panama, but many are rare.

Ocotea cernua

Ocotea cernua

Ocotea cernua (sigua). A small forest tree. The

bark is dark, and there are sometimes small stems sprouting from the base of the trunk. Terminal branchlets are green. Leaves are simple, alternate, shiny green, and have widely spaced secondary veins

and undulate margins. When crushed, they have a musky, avocado-like fragrance, but it is not especially strong. Flowers are tiny, fruits small. Distribution: Widespread in moist and wet zones, from Pacific to Caribbean, always in mature forest. Recognition: The widely spaced and irregular secondary veins help mark this as Lauraceae, and this species is fairly easy to learn from the smallish, undulate leaves. Another Ocotea, O. arcuata (not illustrated) has similar, wavy-margined leaves; it is restricted to wet and montane forest in c. Panama. See also Ocotea whitei. Lacistema aggregatum (Lacistemataceae) has remarkably similar leaves in arrangement and form, but they lack undulate margins, and Lacistema has distinctive scars on the branchlets.

Lauraceae

Nectandra purpurea

Ocotea cernua

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Lauraceae

Ocotea insularis

Ocotea insularis

Ocotea insularis. A medium-sized tree of wet and montane forests. Leaves are simple, alternate,

thick, and fairly tightly clustered toward the ends of branches. There are many secondary veins, but they are vaguely irregular, and crushed leaves smell faintly like avocado. The base of the leaf extends branchward along the petiole. Distribution: Very widespread in lower montane and lowland wet zones. Only in mature forest. Recognition: Easy to miss as Lauraceae if the odor is not checked (or detectable), because the thickened leaves do not look like those of other Ocotea. The extension of the leaf base along the petiole is the best character.

Ocotea whitei

Ocotea whitei

grow in the dry Pacific zone. Many are rare, but there are still quite a few common and widespread species that we have not mentioned.

Persea americana (avocado, aguacate). Not illus-

Ocotea whitei (sigua). A tall old-growth tree. The

trunk is cylindrical and often leans slightly; large trees have substantial buttresses. Some mediumsized trees have stilt-roots, and there are sometimes stems sprouting from the base of the trunk. The bark is dark brown. Leaves are simple, alternate, with a long pointed tip and widely spaced secondary veins. When crushed, leaves smell vaguely like avocado. Fruits are like small avocados with a very thin pulp and a large seed; the fruit base is enclosed in a dark cup. Large adult trees have carpets of seedlings beneath them that last throughout the year. Distribution: Widespread in lower montane and wet forest. Fairly common in the old forest at Barro Colorado I. Not an edge or secondary species. Recognition: This is a difficult species, as the Lauraceae can be. Leaves resemble those of Nectandra purpurea, which has a more curved petiole, O. cernua, which has wavy leaf margins, and O. oblonga (not illustrated). But O. whitei is readily learned as an adult at Barro Colorado I. from the dark bark, big buttresses, and the odd propensity to lean. Most Ocotea species are wet or lower montane species and are found in mature forest, but two species

trated. The yard and garden crop. It can be a fairly large tree. Leaves are large, dark green above, slightly bluish with fine hairs below. Distribution: Widely planted in yards, cities, and farms everywhere in the tropics and subtropics. Probably native to tropical Mexico and mountains of Central America, but so widely planted that its natural range is difficult to decipher. Recognition: This fits Lauraceae in having widely spaced secondary veins and irregularly spaced leaves. The bluish green underside is a common trait of the family as well. Yard trees offer practice in deciphering the Lauraceae leaf odor; it is faint and barely noticeable in older leaves.

Eight species of Persea are native to Panama’s forests. All are rare in Panama but several are more widely seen in Costa Rica’s mountains. The genus also occurs into the s. United States. In most species, leaves are smaller than in the avocado, and there are fine reddish hairs. In addition, there are seven more genera of Lauraceae in Panama we have not mentioned: Aiouea (two species), Aniba (three), Cryptocarya (one), Endlicheria (four), Licaria (six), Pleurothyrium (four), and Rhodostemonodaphne (one). Endlicheria, Licaria, and Rhodostemonodaphne are mostly lowland wet-forest species; the others are largely montane. All are quite rare in Panama and Costa Rica.

Lauraceae

Ocotea insularis

Ocotea whitei

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262

Lecythidaceae

Lecythidaceae Couratari guianensis

Couratari guianensis

The big genus Eschweilera is good to know, but it is difficult.

Couratari guianensis (coquito, condón de mono, zorro, carapelo, congolo). A tall forest tree.

A moderate-sized family of tropical trees, with 292 species worldwide and 204 in tropical America. The Brazil nut, native to Amazonia, is in this family. There are 35 species in Panama, of which we describe 6. The family is best characterized by its fruits, which in most species are hard, woody capsules. In the two biggest genera, the capsules come with tight-fitting woody tops. They hang from branches, and when the top comes loose, seeds fall out. If you are lucky enough to find on the forest floor a capsule plus its matching top (which by this time has separated), you have a cute little wood box with a top. Flowers are large and have many stamens in a characteristic ring. But with leaves and trunk alone, Lecythidaceae are difficult to characterize. The biggest genera have leaves like those of Annonaceae, but sometimes with teeth. Other genera have completely different leaves. A general but not terribly useful character is tough bark, which makes twigs hard to break and bark hard to tear. Learning the Lecythidaceae means learning the species within it.

Trunk straight, cylindrical, with buttresses at the base; bark is gray and flakes off in small pieces. Leaves are simple, alternate, arranged regularly along the branches in a flat plane. Sometimes the leaves are toothed, but sometimes not. The petiole is short and has a slight channel on the upper side. Flowers are pink. The fruits are remarkable: cylindrical woody tubes about 15 cm long with a cap that fits very neatly over the opening at one end. Distribution: In wet forests of c. and w. Panama and nearby Costa Rica. Fairly common on the Caribbean half of the Canal Area. A mature forest tree, not seen in open areas. Recognition: Except where there are fruits on the ground, this species can be difficult to spot. The alternate leaves in a flat plane are typical in other families, and this species might be first identified as an Annonaceae when leaves are lacking teeth. In particular, Guatteria amplifolia has leaves rather like those of Couratari. When leaves are toothed, a first identification might be Salicaceae, but nothing in that family has leaves that are similar. Look especially at the short petioles with a grooved upper surface in Couratari. A second species of Couratari is known only in e.

Couroupita guianensis

Couroupita guianensis

Panama and the Osa Peninsula.

Couroupita guianensis (cannonball tree, bala de cañón, palo del paraíso, coco, palo santo). A big tree

usually known as an ornamental. The trunk is straight and cylindrical. Leaves are simple, alternate, densely bunched toward the ends of branches. Fruits and flowers are produced from the trunk and large

branches, and most trees carry large knobs from past fruiting episodes. The fruits are immense woody balls, hence the common name. Inside there is a dense pulp carrying tiny seeds. Distribution: Occurs naturally in Panama and Costa Rica, but rare and restricted. It is often planted in towns and cities, and Summit Garden has several large trees. Recognition: Ordinarily, you will know this from the fruits themselves or the remnant knobs all over the trunk. As a juvenile, the bunched leaves look like those of species of Pouteria (Sapotaceae), which has latex, or possibly Alseis blackiana (Rubiaceae), though its leaves are opposite. A second species of Couroupita is known from only a few records in far e. Panama and n. Costa Rica. It also has cannonball fruits on the trunk.

Lecythidaceae

Couratari guianensis

Couroupita guianensis

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Lecythidaceae Eschweilera pittieri

Eschweilera pittieri

Eschweilera pittieri (ollito, coquito). Not illustrat-

ed. A medium-sized wet-forest tree. The bark is black, with vertical fissures. Branches emerge perpendicular to the trunk. Leaves are simple, alternate, regularly spaced, and in a flat plane. Secondary veins are

straight, parallel, and impressed above (so raised below). The petiole is dark and has a small channel on the upper side. Flowers are large, whitish yellow, with the many stamens packed into a peculiar thick ball. Fruits are round woody capsules with tightly fitting caps that fall off to free the seeds. On the ground, you find capsules without caps and caps without capsules. Distribution: Common on the Caribbean slope of e. Panama; also on the Pacific slope in w. Panama and the Osa Peninsula. A mature forest tree. This is the only Eschweilera seen in the Canal Area, at Pipeline Rd. Recognition: Leaves look somewhat like those of Guatteria dumetorum (Annonaceae), including the zigzag stem, but are more bullate (impressed veins).

Eschweilera sessilis

Eschweilera sessilis

Eschweilera sessilis (ollito, coquito). A mediumsized tree of cloud forests. The bark is black, with vertical fissures. Leaves are simple, alternate, smooth and thick, almost round, with a prominent yellow main vein but very inconspicuous secondary veins. There are small black dots on the leaf undersides. Flowers are like those of E. pittieri but with pinkish

petals. Fruits are globular woody capsules with a cap. Distribution: A lower montane species from c. to w. Panama, with a few records in Costa Rica. Common at Cerro Jefe near Panama City. Recognition: The leaves do not look like those of other Eschweilera, but have a typical cloud-forest appearance: thick with faint veins. Eschweilera is a large genus, with 14 more species known in Panama. About half of those have highly restricted distributions, and only six of them reach Costa Rica. Those with enough records to tell are from lower montane or wet forest. The typical form is like that of E. pittieri, with distinct and impressed secondary veins. The round woody capsules are always distinctive, but usually are seen on the ground, not the tree.

Gustavia superba

Gustavia superba

Gustavia superba (membrillo). A very abundant,

medium-sized tree. Trunk is straight and only branched near the top. Each big branch has just one cluster of leaves, like a palm. Leaves are very large; no other tree in the area other than palms has larger leaves. They are toothed, long, narrow at the base, and broad at the apex. Flowers are large and purple, with many stamens arranged in a ring, attached to the trunk or large branches just below the leaves. From March to June (at Barro Colorado I.), when they are mature, a few flowers can easily be found on the ground beneath trees. Fruits are round, green, hard, larger and heavier than a softball. Inside are a few large seeds. The fruits remain on the ground for days, rotting and collecting mold. Distribution: A wide-

spread species of lowlands and lower montane sites throughout Panama, including dry parts of the Pacific slope, but not known in Costa Rica. Very abundant in secondary forests. At some sites around the Canal, this species is the dominant secondary forest tree, making up 50% of the canopy. Indeed, its abundance is a good indicator of forest that has been regenerating for 60–100 years. It does not invade during the earliest regeneration, though, the way Cecropia does. Rather, it invades old fields because agoutis carry the seeds into the fields; the very large seeds allow seedlings to germinate in harsh pasture conditions. Recognition: The long leaves in a single cluster at the apex of a large branch or trunk are unmistakable. Other than several rare relatives (see below), just one species is similar, Cespedesia spathulata (Ochnaceae). The two species are unrelated, and have completely different flowers. But without flowers they are easily confused. With a fallen leaf in hand, look closely at the leaf underside—the smallest veins of Cespedesia are prominent and parallel, whereas in Gustavia they are inconspicuous and widely spaced. On the trees, leaves of Gustavia tend to droop downward much more than leaves of Cespedesia, some of which point upward, and new leaves of Cespedesia

Lecythidaceae

265

Eschweilera sessilis

Gustavia superba

are conspicuously red, unlike those of Gustavia. A species called Clavija costaricana (not illustrated; in the family Theophrastaceae) has leaves in the same pattern as Gustavia, but Clavija is a little treelet, never more than a couple meters tall.

Eight more Gustavia species are found in Panama, but they are sparse and seldom seen, and only one of those reaches Costa Rica. Some closely resemble G. superba. But others, such as G. dubia, have much smaller leaves.

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Lepidobotryaceae Lecythis ampla

Lecythis ampla

Lecythis ampla (olla de mono, coco). A wet-forest

giant. The trunk is straight and tall, unbranched until high. The gray bark has deep vertical fissures. Leaves are small, simple, alternate, finely toothed, regularly spaced along the branches, and in a flat plane. Flowers are large, pink, with hundreds of stamens packed in a ring. The fruit is an enormous and very heavy

woody capsule with a top that comes free. Distribution: Known in wet forest on the Caribbean slope, but sparse and not common. Recognition: Fruits on the ground are unmistakable, and the huge fissured trunk is a good clue. Toothed, alternate leaves should recall Salicaceae, but this looks more like Humiriastrum diguense (Humiriaceae). Four more Lecythis species are known in Panama, three of which are restricted to the eastern half of the country; the last is sparse in Panama and Costa Rica. The only genus of Lecythidaceae we have not yet mentioned is Grias. The one species, G. cauliflora, has leaves very much like those of Gustavia superba, but they lack teeth and have short petioles (almost none at all).

Lepidobotryaceae Ruptiliocarpon caracolito

Ruptiliocarpon caracolito

A tiny family of tropical trees with a peculiar distribution: a single species in Africa and a single species in South America. With just the one species in Panama, we omit a family description.

Ruptiliocarpon caracolito. A tall wet-forest

tree. The trunk is straight, cylindrical, and bark reddish with big warts. Leaves are small, simple, alter-

nate, shiny green, and smooth. The petiole is green and divided into two sections by a ring around the middle. Fruits are green capsules that open to reveal aril-covered seeds. Distribution: Only in wet and lower montane forests, where widely distributed in Costa Rica; in Panama principally in the Canal Area. Recognition: The leaves are not by themselves distinct, broadly resembling a range of other small, alternate-leaved trees (Drypetes [Putranjivaceae], Margaritaria [Phyllanthaceae], and Oxandra [Annonaceae]), but note the unusual and distinct petiole. It resembles a Fabaceae petiole, and so Ruptiliocarpon could be confused with those unusual Fabaceae with simple leaves, such as Swartzia or Dalbergia monetaria (both Papilionoideae); the latter has especially similar leaves, but we do not illustrate it.

Lepidobotryaceae

Lecythis ampla

Ruptiliocarpon caracolito

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268

Lythraceae

Lythraceae Lafoensia punicifolia

Lafoensia punicifolia

A moderate-sized family of aquatics, herbs, shrubs, and trees, found in both tropical and temperate zones. There are 600 species worldwide and almost 400 in tropical America, but few of those are trees. This is the pomegranate (Punica granatum) family, but that is an Asian species. In North America, purple loosestrife (Lythrum), which grows widely in marshes, is in this family. Panama has only two native trees, plus a widely planted ornamental from Asia. We describe just one species. Lythraceae is one of the few families with opposite leaves, but there are so few species that are trees it does not pay to learn family-level traits.

Lafoensia punicifolia (amarillo, amarillo pepita, calabacito). A large, sparsely distributed tree. Large

trees have small buttresses at the base. The bark is gray, with vertical fissures. Leaves are small, opposite, simple, regularly spaced, and in a flat plane; the

small size and regular spacing suggest a compound leaf with many small leaflets. Each leaf has an extended but blunt tip at its apex, and underside this tip is a tiny brown cavity (called a domatium). Some leaves turn yellow during the dry season. Flowers are large, bell-shaped, yellowish or yellow-green, at the ends of the branches. Fruits are woody capsules that open during the dry season to release winged seeds. Distribution: Known at scattered sites, especially on the Pacific slope. Recognition: The opposite and regularly spaced leaves look like those of Myrtaceae, for instance Eugenia nesiotica, but see other Myrtaceae as well. Myrtaceae usually have translucent spots in the leaf. Mouriri myrtilloides (Melastomataceae) also has leaves similar to those of Lafoensia, but it lacks petioles. None of these other species reaches large size, and none has a domatium at the leaf tip. You might think leaves of Lafoensia are compound, but look carefully at the base of the branch and the number of leaves per branch. The other native tree in the Lythraceae is Adenaria floribunda. It is a small, weedy shrub with opposite leaves. An Asian species, Lagerstroemia speciosa, known as crape myrtle or reina de las flores, is planted as an ornamental, abundantly so along streets of the Canal Area. It has opposite leaves and big purple flowers.

Magnolia sororum

Magnoliaceae Magnolia sororum

A modest family of 220 species of trees worldwide, most occurring from China south into the Asian tropics. There are several species in eastern North America, and 60 in tropical America. The genus Magnolia is a well-known temperate tree, often planted as an ornamental for its big, showy flowers. There are five species in Panama, and we cover one. Magnoliaceae have alternate leaves and circular scars on the branches at the base of leaves. Crushed leaves, and especially unopened buds, have a strong, spicy odor. Flowers are large and showy. Fruits are covered in scales and vaguely resemble pinecones, though they are not woody. The circular twig scars look like those of Moraceae, but Magnoliaceae lack latex and have much different flowers.

Magnolia sororum (baco). A tall, montane tree.

Twigs have distinct encircling scars at the base of each leaf, where the stipule has fallen. There are fine, silvery hairs on the twigs. Leaves are simple, alternate, bunched toward the ends of branches, glossy green above but paler below. Flowers are white and large. Fruits are scaly capsules that open to reveal red seeds. Distribution: Known from Cerro Campana, w. Panama, and c. Costa Rica, at mid- to high elevation. Recognition: If you know the flowering Magnolia of the s. United States, these leaves will look familiar. The stipule scar resembles those of Ficus and other Moraceae, but all Moraceae have lots of white latex and Magnolia has none. Two or three more Magnolia species are found in Panama, all in montane forest. One is endemic to Panama, whereas another is also found in c. Costa Rica. The genus is mixed up with Talauma, the other member of the Magnoliacaeae known in the area, and there are four or five species total, some going by both names. Talauma is widespread in lowland wet forests in Panama and Costa Rica, but also known in mountains. Like Magnolia, it has showy, white flowers, and shiny leaves with stipule scars.

Magnoliaceae

Lafoensia punicifolia

Magnolia sororum

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270

Malpighiaceae

Malpighiaceae Bunchosia nitida

Bunchosia nitida

This is a large family of strictly tropical trees and lianas, with 1260 species worldwide and 1110 in the Americas. None of its species are widely known, but many have edible fruits, including the acerola, or Barbados cherry (Malpighia glabra). In Panama there are 24 species of trees and shrubs (plus many more lianas), of which we describe 5 and illustrate 4. This is one of the minor families with opposite leaves. It might be confused with Myrtaceae, but lacks the collecting vein. It might be confused with Rubiaceae, but stipules in Malpighiaceae sit in the leaf axil, not between leaves as in Rubiaceae (intrapetiolar, not interpetiolar). A minute character found throughout the Malpighiaceae is T-shaped hairs; these require magnification to see. In some species, hairs are sparse and perhaps only found on young twigs. Flowers are consistent across the family; usu-

ally yellow or orange, with petals that widen above like little claws.

Bunchosia nitida (cerezo de monte). A mediumsized tree of the Pacific coast. Trunk is straight but fluted. Leaves are simple, opposite, spaced regularly along the branch, in a flat plane. The leaf underside has silvery pubescence creating a reflective sheen. There may be small glands near the base on the leaf underside: elevated disks of yellow or green. The flowers are bright yellow, and fruits are fleshy, single seeded. Distribution: Mostly near the Pacific coast throughout, and shows up commonly on rocky limestone (Madden Dam, for example). In the Canal Area, widespread but sparse from Panama City to Gamboa, where found along roadsides. Recognition: Quite similar to another species in a minor, opposite-leaved family, Aegiphila panamensis (Lamiaceae), which also has a silvery sheen below. Glands of Bunchosia provide a distinction, but they are subtle. Eight more Bunchosia species are known in Panama. B. odorata is from dry sites near the Pacific, and three others are from wet or lower montane forest; the remaining are rarely seen. Most have green, not silvery, leaf undersides, and most could be confused with Aegiphila species (Lamiaceae).

Byrsonima crassifolia

Byrsonima crassifolia

Byrsonima crassifolia (nance). A farmland tree. The trunk often has low branches, so the crown is close to the ground, but it can grow fairly tall and straight. Leaves are opposite, tightly clustered at the end of branches, dark shiny green on the top, dense-

ly pubescent on the underside. The flowers are yellow-orange, produced in clusters above the leaves; the petals are narrow near the base, then broaden. Fruits are round berries, green initially, maturing yellow, edible. Distribution: Abundant in farmland and residential areas throughout the lowlands, never in any sort of woods or forest. Resistant to fire, and common in burned grasslands of the dry rural zone. Recognition: Most easily known by its occurrence around human settlements, where the crowded, opposite leaves with densely hairy undersides distinguish it. Flowers are conspicuous and distinctive, and the fruit is very well known, sold on the street, and usually made into a drink.

Malpighiaceae

Bunchosia nitida

Byrsonima crassifolia

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Malvaceae

Byrsonima spicata

Byrsonima spicata

Byrsonima spicata (nancito, nancillo, nance). A

tall forest tree. The trunk is usually straight and cylindrical, and the bark dark gray. Leaves are simple, opposite, narrow and pointed at the tip, with a slightly wavy margin. They are crowded toward the end of branches, and each successive leaf pair is perpendicular to the previous pair. Leaves turn yellow as they

age, and large crowns usually have a few yellow leaves. The petiole is short and covered in fine hairs, and the small, triangular stipule is also hairy. Flowers and fruits resemble those of B. crassifolia. Distribution: Known only in c. Panama, where found throughout Canal Area forests, from Pacific to Caribbean. Never common and not well known; not seen in farmland or along roads. Recognition: The narrow leaves are much different from those of B. crassifolia, so the two are not confused. Panama has three more species of Byrsonima, all forest trees and none anywhere near as common as B. crassifolia. A wet-forest species, B. crispa, is common in Costa Rica but rare in Panama. The other two species have very restricted, montane distributions.

Spachea correae

Spachea correae

in Panama. Recognition: The glossy thick leaves are unlike other Malpighiaceae. Rudgea (Rubiaceae) has similar leaves, but with different stipules and no latex. S. correae might be confused with Apocynaceae (opposite leaves and latex), but glands and stipules are different. It might also recall Myrtaceae, but no Myrtaceae has glands or latex.

Spachea membranacea. A rare forest tree. Spachea correae. Not illustrated. A rare forest

tree. The trunk is straight, with black bark. Leaves are simple, opposite, held perpendicular to the branch (do not lie flat), and are wide, with round tips, very thick, and with faint venation. The top side is glossy green, and the underside lighter with (usually) some tiny, yellow, disk-shaped glands. Broken leaves produce white latex that appears slowly. Flowers are pink to purple, held on a spike, with narrow, cupped petals. Fruits are small berries. Distribution: Known on the Caribbean coast from the Canal through Costa Rica, plus the Osa Peninsula. Very rare

Malvaceae

Leaves are simple, opposite, shiny green above and below. There is a pair of light-colored, circular glands at the very base of each leaf (on each side of the petiole). Broken leaves exude white latex. Flowers are pink. Distribution: A few individuals are known on Barro Colorado I.; only known from two other sites. Recognition: Like S. correae but leaves are not pointed at the tip. Should suggest Apocynaceae and Myrtaceae. Spachea is so rarely seen, remembering to look for the little glands is a challenge. One other Spachea species is known in Panama, and it is also very rare.

Spachea membranacea

(Including Bombacaceae, Sterculiaceae, Tiliaceae, and Malvaceae) Spachea membranacea

These four major plant families of the tropics were always known to be related. Recent analyses, including DNA sequences, now show them to be intermingled

with one another, and the four are now treated in one much larger group that takes the name Malvaceae. Here we treat the wide family together, but follow an arrangement of subfamilies that helps separate the groups within. One trait characterizes them all: palmate venation at the leaf base, that is, three to five major leaf veins arising where the petiole meets the leaf. Moreover, many species in the group have palmately lobed or palmately compound leaves. In this broad Malvaceae, the Panama checklist has 92 tree or treelet species, of which we cover 23 and illustrate 22.

Malvaceae

Byrsonima spicata

Spachea membranacea

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Malvaceae—Bombacoideae Cavanillesia platanifolia

Cavanillesia platanifolia

Bo MBac o IDeae (subfamily within Malvaceae). Now relegated to a subfamily, this is

a medium-sized group as far as species number goes, from both the American and the Old World tropics, but it includes some of the biggest trees in the world. There are 35 species of the Bombacoideae in Panama, and we cover 8. The best-known Bombacoideae have enormous, bloated trunks or huge plank buttresses, like the kapok tree (Ceiba) of South America and Africa. But not all are big trees, and there is a major group in South America with many slender treelets. The first group is easier to recognize, because all have palmately lobed or compound leaves, but the second group is difficult. It includes many treelets plus some tall trees, and all have simple, alternate, unlobed leaves carried on a petiole swollen at both ends but narrow in between. They have palmate venation, as in the rest of the Malvaceae, and generally have verticillate branching, meaning groups of branches emerging from a single spot on the main trunk. The two groups of bombacoids differ ecologically: the first comprises fast-growing trees of open areas, whereas the second inhabits mature forest understory. The first group—the giant trees—has more genera and is better known, but the second group has more species. The eight species we cover include seven from the first group and just one from the second.

Cavanillesia platanifolia (cuipo). An unmistakable giant with a bizarre trunk. Large individuals have straight trunks, without buttresses, but are bloated at the base. The bark is very smooth, except for horizontal rings every few meters. The trunk is unbranched for most of its height. The crown has a flat base and a round top. Leaves are large, almost round on adults, but on juveniles are palmate and almost square. This is one of the truly deciduous species in the area, dropping all its leaves in November, before the rains stop, and regrowing them in April or May, after rains begin. It produces pinkish orange flowers and then fruits when leafless. The fruits consist of five large wings, all semicircular, nearly at right angles to one another. Distribution: A species of the drier regions of c. Panama, but it also appears on limestone or other rocky soils in wet areas. It frequently associates with Bursera simaruba (Burseraceae) and Calycophyllum candidissimum (Rubiaceae). Common in forests of the drier, Pacific half of the Canal Area, and occasionally in agricultural areas away from forest. Saplings are scarce: its populations are nearly always just a few large to giant trees, but it only takes a few cuipo crowns to fill a forest. Recognition: Nothing like it when larger than about 10 cm in stem diameter. When leafless in the dry season, the giant crown is conspicuous from kilometers away, even more so when it fruits. When studying leaves on smaller trees, it might be confused with the balsa (Ochroma), which also has large, squarish leaves, but the trunk is completely different. Sterculia apetala (Byttnerioideae) is another large tree with big leaves and a geographic range similar to that of cuipo (for example, common around Panama City). See also Heliocarpus americanus (Grewioideae).

Ceiba pentandra

Ceiba pentandra

Ceiba pentandra (kapok, ceibo). One of the largest trees of the area, with immense plank buttresses. In big trees, the trunk above buttresses is straight and cylindrical, with gray bark. There is generally one set of enormous horizontal branches at the top—the trunk ends where it branches. The crown is as wide as any in the world, but has sparse foliage. The leaves are palmately compound, with narrow, pointed leaflets arranged radially; secondary veins are not prominent. Saplings have sharp spines all over the trunk, plus vertical green lines in the bark. In some years, all leaves fall in December, but readily regrow, and

no flowers are produced. In other years (or in other trees), leaves fall and flowers are produced in December, then trees remain leafless for several months while fruits develop. Flowers are white to various shades of off-white. The seeds break out of a dry green pod and are covered by cottonlike filaments (the kapok), so they float in the wind. Distribution: There are occasional giant trees in most forested areas and sometimes pasturelands throughout the lowlands. Seldom seen as a juvenile. Recognition: Large trees should be unmistakable, because other species with large buttresses (Ficus insipida [Moraceae] and Sterculia apetala [Byttnerioideae]) do not have such fat trunks or gray bark. The palmately compound Bombacoideae (Ceiba, Pachira, and Pseudobombax), plus Jacaratia (Caricaceae) and the crop plant manioc (Manihot [Euphorbiaceae]), all have similar leaves; see the discussion under Pseudobombax. Ceiba contains one other species, which is seldom seen.

Malvaceae—Bombacoideae

Cavanillesia platanifolia

Ceiba pentandra

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Malvaceae—Bombacoideae Ochroma pyramidale

Ochroma pyramidale

Ochroma pyramidale (balsa, balso). A mediumsized tree of roadsides. The trunk can be straight, but often forks near the ground; has only small buttresses. Leaves are simple, alternate, very large, on juveniles the size of a small umbrella. They are slightly lobed, almost square, with three to five prominent veins arising from the base and a whitish underside. Flowers are very large, to 20 cm across, white, produced in December and January and very attractive to many nectar-feeding birds and mammals. Fruits

are green capsules enclosing seeds that carry cottony filaments like kapok. Distribution: Grows in nearly any open area from lowlands to low mountains, and is abundant along roads. Inside the forest, it is only found in big clearings. Juveniles are seldom seen. It is one of the fastest growing trees in the world, and large trees show up a few years after any sort of forest-clearing operation. Recognition: Will be readily learned because of its abundance and very large leaves. The nonnative teak, Tectona grandis (Lamiaceae), has leaves with nearly the same form, but opposite, not alternate. Cecropia (Urticaceae) also has large leaves, but very different in shape. See also Cavanillesia of this subfamily, plus Croton draco and Jatropha (Euphorbiaceae), also with large, wide leaves. This is the balsa of model-airplane fame, with extremely light, soft wood. It is also used for making rafts (hence the name balsa, Spanish for raft).

Pachira aquatica

Pachira aquatica

Pachira aquatica (sapote, coco de agua). A medi-

um-sized forest tree of swamps and mountains. Trunk is straight, with small buttresses. Leaves are alternate, palmately compound, without teeth. Leaflets have a small point at the tip, and secondary veins are

prominent. Flowers are very large, with long, narrow, pinkish petals and dozens of long stamens in a shaving-brush form. Fruits are capsules, but seeds are large and woody and have a fleshy pulp. Distribution: In two different habitats: lowland swamps along the Caribbean coast plus the Osa Peninsula, where it seems to be more common, and secondarily in lower montane cloud forest. Recognition: The palmately compound Bombacaceae (Ceiba, Pachira, and Pseudobombax), plus Jacaratia (Caricaceae) and manioc (Manihot [Euphorbiaceae]), all have similar leaves; see the discussion under Pseudobombax. This species often has fruits on it, and they are not at all like those of the other Pachira, all of which have cottony kapok on tiny seeds.

Malvaceae—Bombacoideae

Ochroma pyramidale

Pachira aquatica

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Malvaceae—Bombacoideae Pachira quinata

Pachira quinata

Pachira quinata (cedro espino). A tall tree with a

large trunk and modest buttresses. Small and medium-sized individuals have sharp spines on the trunk, slightly curved at the tip; in large trees, spines can be sparse, but there are usually at least some present. Leaves are alternate, palmately compound; leaflets have small teeth and are either blunt or with a small point at the tip; secondary veins are prominent. Completely leafless during the dry season, when the large

flowers are produced, like those of P. aquatica but white. The fruit is a dry capsule that splits to release tiny seeds on cottonlike hairs. Distribution: Mostly a species of the drier Pacific slope, but not common. Also shows up on limestone in wetter areas. Like the other big bombacoids, it is only encountered as a large tree, not as a sapling. This is one of the most widely planted native species in plantations, because the wood is valuable. Recognition: Spines on the trunk of big trees are a clear character, but see also Hura crepitans (Euphorbiaceae), which lacks buttresses, and Zanthoxylum (Rutaceae), spines of which are much different. As a juvenile, Ceiba pentandra also has spines and has similar leaves. The palmately compound Bombacoideae (Ceiba, Pachira, and Pseudobombax), plus Jacaratia (Caricaceae) and manioc (Manihot [Euphorbiaceae]), all have similar leaves; see the discussion under Pseudobombax.

Pachira sessilis

Pachira sessilis

Pachira sessilis (yuco de monte, ceibo). A tall forest tree with a straight and cylindrical trunk. Large individuals have flat, thin buttresses at the base. The bark is yellowish, lacks spines, has vertical fissures,

and sometimes has small pieces peeling free. Leaves are alternate, palmately compound, without teeth, blunt at the tip; secondary veins are not prominent. Becomes leafless during the dry season. Flowers are like those of the other Pachira, but darker in color; fruits are capsules with tiny, kapok-laden seeds. Distribution: Mostly along the Pacific coast, including fairly dry forests, but also found at wet and lower montane sites. Saplings mostly restricted to forest clearings or open areas, and not seen often. Large trees are occasionally seen along roads throughout the area. Recognition: See the similar Pachira quinata and Pseudobombax.

Malvaceae—Bombacoideae

Pachira quinata

Pachira sessilis

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Malvaceae—Bombacoideae Pseudobombax septenatum

Pseudobombax septenatum

Pseudobombax septenatum (barrigón). A large tree of open areas or forest clearnings. Trunk is tall and straight, lacks buttresses, but is swollen at the base. The bark has vertical green lines (the green is chlorophyll!). Leaves are alternate, palmately compound. Leaflets lack teeth and have a weakly pointed tip; secondary veins are prominent. Trees becomes completely leafless in the dry season, and then produces big, white, pom-pom–like flowers (like shaving brushes, if the cheerleading analogy doesn’t work). Fruits are dry capsules carrying tiny seeds with cottony filaments. Distribution: Widespread in secondary forests and open areas, especially on the drier Pacific slope. Rare in mature forest where it is only found in large clearings. Saplings are scarce.

Recognition: We have just covered five bombacoids that are large trees with fairly similar palmately compound leaves. Of the group, only Ceiba has narrow leaflets with long, pointed tips. Both Ceiba and Pachira sessilis have faint secondary veins, but the latter has blunt leaflets. The remaining three, Pachira aquatica, Pachira quinata, and Pseudobombax septenatum, are similar to one another. Moreover, in very big trees, details of leaf form cannot be compared, so one must study the trunks. In big trees, only Pseudobombax has vertical green lines in the bark (in saplings, Ceiba does also). Only Ceiba and Pachira quinata have spiny trunks, and Ceiba loses its spines when it is big. As large trees, Ceiba and Pachira sessilis are most likely to be confused, because they have similar, spineless bark; if you can see the leaflets, those of Ceiba are pointed and those of Pachira are blunt. Then there is Jacaratia spinosa (Caricaceae), completely unreleated but with palmately compound leaves and a spiny trunk; when small, it gets confused with Ceiba, but it is never really big, and has no buttresses. Finally, the crop manioc (Manihot [Euphorbiaceae], not pictured) has leaves like those of Ceiba, but lacks spines and grows like a shrub in farms and gardens.

Quararibea asterolepis

Quararibea asterolepis

Quararibea asterolepis (guayabillo). A tall, old-

forest species unlike the previous seven Bombacoideae. It has a straight trunk branching only near the top. At the base, the trunk is fluted or flanged, not cylindrical and sometimes nearly square in cross section. At the base, the the flanges usually spread to form narrow but tall buttresses. The bark peels off, leaving a smooth, naked, light gray trunk, with irregular vertical greenish or dark gray bands. Branches emerge perpendicular to the trunk, four from one spot (verticillate branching); each set of horizontal branches can form a distinct layer, separate from the next layer up. Leaves are alternate, oval, with three major veins emerging from the base. The petiole is swollen just below the leaf, but narrow below the swelling. Flowers are moderate-sized, cream-colored. Fruits are spherical, with large seeds and a sweet pulp within. Distribution: Widespread on the Caribbean slope of Panama and in wet forests throughout Costa Rica; occasionally in lower montane forest (Cerro Campana near the Canal). One of the dominant canopy trees in Barro Colorado I. old growth,

and saplings are abundant in the forest shade there. Never seen in farmland or residential areas. Recognition: Unlike and never confused with the other Bombacoideae. Medium-sized and large trees are readily recognized by the peculiar, smooth, flanged trunk. Guapira standleyana (Nyctaginaceae) has a ribbed trunk of the same size and shape that of Q. asterolepis, but it has brown bark, not like the smooth and naked trunk of Quararibea. Small Quararibea lacking the trunk characters are easily overlooked. Note the verticillate branching, palmate venation, and swollen petiole. Several Theobroma species (Byttnerioideae) closely resembly Quararibea in leaf form, but lack verticillate branching. Ten other Quararibea species are included in the Panama checklist. Seven of those are very rare, and the other three are restricted to wet forests. In addition, there are 12 species in the similar and related genus Matisia (none of which is illustrated), but most of those are rare and some have very restricted distributions. They share palmate venation, swollen petioles, and verticillate branching; one species, M. obliquifolia, is common on the Caribbean slope in both Panama and Costa Rica and has very large, asymmetric leaves that are easy to spot. Four other genera of Bombacoideae are found in Panama, each with a single species. Bernoullia and Gyranthera have compound leaves much like those of Pseudobombax, whereas Patinoa and Phragmotheca are closely related to and resemble Quararibea.

Malvaceae—Bombacoidea

Pseudobombax septenatum

Quararibea asterolepis

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Malvaceae—Byttnerioideae Guazuma ulmifolia

Guazuma ulmifolia

Byttne RIo IDeae (subfamily within Malvaceae). This subfamily includes many of the

genera formerly classified in the family Sterculiaceae; see the Sterculioideae for the rest. They are mostly treelets or small trees, and include perhaps the most prized tree in the world, chocolate (Theobroma cacao). One group has palmately compound leaves (Herrania), but others have simple leaves, though always with palmate leaf venation. All told, the Panama checklist has 13 species of trees and treelets in the Byttnerioideae, and we cover 4.

Guazuma ulmifolia (guácimo). A spreading, medium-sized tree of farms and roadsides. The trunk usually leans and has branches near the ground,

though rare forest individuals grow straight and tall. Leaves are alternate, in a flat plane along long branches. The leaf base is very asymmetric, and leaves are rough (asperous) to the touch. There are three main leaf veins arising together from the leaf base. Flowers are small, whitish cream in color. The fruits are woody and reminscent of tiny pine or alder (Alnus [Betulaceae]) cones. They are green when immature, but turn black, and that is how they are found in great numbers beneath trees. They do not look appealing, but have a cherry flavor if you are willing to try chewing one. Distribution: A very common tree of farmland and roadsides, sometimes forming nearly pure stands in abandoned pasture. Fairly common inside secondary forest, and, remarkably, it turns up as a few large trees in the old forest on Barro Colorado I. Recognition: Not difficult to learn because of its abundance in residential areas, and the asymmetric leaf base and asperous feel are distinctive. Trichospermum (Malvaceae—Grewioideae), Muntingia (Muntingiaceae), and Trema (Cannabaceae) all have fairly similar leaves, and Muntingia also grows in pasture and abandoned farmland. None of those have asperous leaves, and their fruits are much different.

Herrania nycterodendron

Herrania nycterodendron

Herrania nycterodendron (cacao de monte). A

wet-forest treelet. It is a single, straight, thin stem with a few compound leaves at the top. Each leaf has five large, toothed leaflets, and the leaf as a whole is held

on a long petiole. Flowers and fruits are produced on the trunk well below the leaves. Flowers are deep red and stalkless. The fruit resembles that of the cacao (chocolate), a ribbed pod that turns from green to yellow as it matures. Distribution: We mistakenly called this H. pulcherrima for several years, but in 2003 determined that it was in fact H. nycterodendron, a species not then known in Panama. Still only known from a few of our inventories on the Caribbean side of the Canal Area. Recognition: Distinguished from H. purpurea by toothed leaflets. Might be confused as a sapling with Ceiba, Pachira, or Pseudobombax (Bombacoideae), but their leaves lack teeth.

Malvaceae—Byttnerioideae

Guazuma ulmifolia

Herrania nycterodendron

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Malvaceae—Byttnerioideae Herrania purpurea

Herrania purpurea

Herrania purpurea (cacao de monte). A small

forest treelet just like the previous except having untoothed leaflets. The seeds (inside the pod) have a sweet, white pulp that is tasty. Distribution: In the interior of mature, lowland forests of moist to dry sites. Quite common at Barro Colorado I. Recognition: Except for the previous species, there is nothing like it in the forest shade. Leaves are reminiscent of those of Pseudobombax and Pachira (Bombacoideae), but saplings of those species are rare inside the forest. The one other Herrania species in the Panama checklist is H. pulcherrima, the one we mistook for H. nycterodendron. A few specimens are known from the Caribbean coast in e. Panama, but it seems likely they are really H. nycterodendron.

Theobroma cacao (cacao, chocolate). An unpresumptious treelet of forest shade whose spindly ap-

pearance seems an unlikely source of its ecomonic and cultural power. Has a weak, sometimes leaning stem, branching near the ground. Leaves are simple, alternate, long and rather narrow. There are three large veins palmate at the base, and the petiole has a swelling at both ends. Flowers and fruits are produced on the trunk, sometimes not far from the ground. The wrinkled, ridged pod starts green and turns yellow, and has many black seeds within that are the source of its value. The white pulp around the seeds is sweet and tasty. Distribution: The cacao was originally Amazonian, but the Mayans brought it to Mexico centuries ago. It is cultivated now along the Caribbean coast in both Panama and Costa Rica, and spreads from plantations into nearby forests. There are a few in the old forest at Barro Colorado I. Recognition: Cacao is easy to overlook, because the leaves are not striking and fruits are not often present. The palmate venation and swollen petiole are good characters, but see Quararibea (Bombacoideae), which has remarkably similar leaves. Panama has six species of native Theobroma, all understory treelets resembling the cacao. T. bernouillii has long, narrow leaves with very widely spaced secondary veins and grayish white leaf undersides; it is common in the Fort Sherman area near the Canal and elsewhere along the Caribbean coast of Panama (but not Costa Rica). Others are restricted to wet or montane forest.

Malvaceae—Byttnerioideae

Herrania purpurea

Theobroma cacao

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Malvaceae—Grewioideae Apeiba membranacea

Apeiba membranacea

GRew Io IDeae (subfamily within Malvaceae). This subfamily includes most of what used to

be the Tiliaceae, but see the Tilioideae for one other. It is a worldwide group of tropical trees, few in species, but among them are some of the most abundant trees of roadsides and clearings of Panama and Costa Rica. All grewioids have simple, alternate, toothed leaves with well-defined palmate venation. These are solid characters; there are other genera and families with the combination, but not many. There are 13 trees in the Panama checklist; we cover 6 of them.

Apeiba membranacea (peinecillo, cortezo,

monkey comb). A tall tree of wet to moist forest. The

trunk is slightly ribbed or irregular, weakly swollen at the base to form small buttresses. Bark is light gray. Leaves are simple, alternate, sometimes slightly toothed, arranged regularly along branches, in a flat plane, and light gray on the underside. There are three main veins arising together from the leaf base, and tertiary veins are dense and parallel. The petiole is swollen where it meets the leaf. Flowers are yellow, produced above the crown. Fruits look just like black sea urchins, with dense, woody spines covering a black capsule. Distribution: Widespread but never especially common in moist and wet forests, not outside the forest and only occasionally along forested roads. Saplings are scarce and restricted to natural forest clearings, so you will mostly encounter big trees in the forest. Recognition: The three prominent leaf veins and dense ladderlike tertiary veins are distinctive. Only the other Apeiba and the two Luehea are similar. But of those, only L. speciosa might be confused with A. membranacea, because it also has a light leaf underside, though whiter than in A. membranacea. Inside the forest, where saplings are scarce, you can learn to recognize the grayish trunk; look for both leaves and fruits on the ground.

Apeiba tibourbou

Apeiba tibourbou

Apeiba tibourbou (peine de mono, peinecillo, cortezo). A medium-sized tree of open lands. It is often forked near the ground and has long, hanging branches. Occasionally in the forest it grows straight, but not as tall as the canopy. Twigs have dense, red hairs. Leaves are large, alternate, finely toothed, arranged in a flat plane and spaced regularly along the branches, rough to the touch. Unlike the previous species, they are green below. There are three main veins arising together from the leaf base, and tertiary

veins are closely spaced and ladderlike; veins are raised and conspicuous on the underside of the leaf. Flowers are yellow, flat, star-shaped, with pointed petals, and present throughout the wet season. Fruits are dry capsules densely covered with soft green spines. Distribution: An abundant tree of abandoned farms, roadsides, and grasslands, especially along the dry Pacific slope. Sometimes forms dense bushy stands, and if grass is prevented from burning, this species colonizes quickly. Very rare in mature forest; only in natural clearings. Recognition: Other roadside species with leaves of similar size and shape are Castilla elastica (Moraceae) and Annona spraguei (Annonaceae); both lack palmate venation and are not asperous. In the forest, leaves of Quararibea asterolepis (Bombacoideae) are similar but not toothed. In a pinch, look for the dense, red hairs on twigs of Apeiba. The previous species, A. membranacea, has quite different leaves and is not confused with A. tibourbou.

Malvaceae—Grewioideae

Apeiba membranacea

Apeiba tibourbou

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Malvaceae—Grewioideae Heliocarpus americanus

Heliocarpus americanus

Heliocarpus americanus (majaguillo, majagua).

A medium-sized tree of cloud forest. It has a straight, cylindrical trunk, often branched near the base. Leaves are simple, alternate, toothed, on long, pubescent petioles, heart-shaped, sometimes with three pointed lobes. Three or five main leaf veins arise from the leaf base, and notice the tiny glands near

the leaf base. Flowers are yellow, in dense terminal clusters. Seeds are very peculiar: pinkish feathery tufts in dense clusters, they look like bryozoans or some other coral reef creature. Distribution: Mostly in lower montane forest, at a few scattered sites and not common. Known in lowlands of the Canal Area too, where it can be seen occasional in clearings around Panama City. Most often seen in clearings and disturbed areas in the cloud forest at Cerro Campana. Recognition: Could be overlooked as a balsa (Ochroma in the Bombacoideae), which has leaves of the same form, or the cuipo (Cavanillesia, also Bombacoideae), which has large, lobed leaves, but a much different trunk. See also the genus Triumfetta, mentioned at the end of the section on Grewioideae but not illustrated.

Luehea seemannii

Luehea seemannii

Luehea seemannii (guácimo colorado, guácimo

molenillo, guácimo pacheco). A tall tree of roadside and edge. Large trunks are somewhat fluted or have irregular buttresses; bark is light gray or brown. Branching is irregular, sometimes near the ground, sometimes not. Leaves are alternate, regularly spaced along branches, and have three main veins emerging from the leaf base; tertiary veins are parallel and closely spaced. Underside of the leaf is conspicuously brown, due to fine brown hairs, and this is

easy to see in even the largest canopy individuals. In very small saplings, the underside can be whitish rather than brown. Produces large numbers of whitish flowers in December and January, with tiny petals, looking like a ball of many stamens. A number of bird species, such as Tennessee warblers and orchard orioles, visit the flowers for nectar. Fruits are small, ribbed capsules. Distribution: Numerous along nearly every road and forest edge, also in farmland. One of the dominant species of secondary forests in all areas on the Pacific half of the Canal Area. Also occurs sparsely in old-growth forest, where it can be a tall tree with a wide crown. Inside the forest, it is rare as a sapling, only appearing in natural tree-fall clearings. Recognition: Unmistakable. Other species with conspicuously brown leaf undersides (Chrysophyllum cainito [Sapotaceae] and Vismia macrophylla [Hypericaceae]) have very different leaves. See also L. speciosa, with whitish leaf undersides.

Malvaceae—Grewioideae

Heliocarpus americanus

Luehea seemannii

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Malvaceae—Grewioideae Luehea speciosa

Luehea speciosa

Luehea speciosa (guácimo molenillo, guácimo blanco, guácimo borcico, guácimo tortugo). A medium-sized tree of roadside and edge. Leaf shape and arrangement are just like those of L. seemannii, but the underside is conspicuously white, not brown, due to fine white hairs. Also differs from L. seemannii in

having large, white, Hibiscus-like flowers, with just few flowers on a tree. Distribution: Same roadside habitat as L. seemannii, but mostly on the Pacific slope in dry c. Panama and nw. Costa Rica. Not especially common, nothing like the abundant L. seemannii. Recognition: The white leaf undersides with palmate venation are distinctive, unlike any other tree of the area. In December, the large white flowers are easy to spot, even from a distance, once they are learned. One more species of Luehea is found in Panama, L. candida. It is similar to L. speciosa, but with wider leaves, and occurs only in the driest parts of Panama and Costa Rica.

Trichospermum galeottii

Trichospermum galeottii

Trichospermum galeottii (capulín, majaguillo, burrilico). A medium-sized forest-edge tree. Leaves are alternate, simple, with fine teeth, long and straight-sided (unlike more heart-shaped leaves of its relatives); three major veins arise at each leaf base and then turn parallel to the leaf edge. It produces conspicuous purple flowers in November (in the Canal Area) when few other trees are flowering. Distribution: Frequent in the Canal Area; else-

where, known at a few scattered locations. Mostly in the lowlands, both Caribbean and Pacific slopes, but also at lower montane sites. Sometimes in yards or farms, and also inside the forest, though only in natural gaps. Saplings are seldom seen. Recognition: Purple flowers on roadside trees at the right time of year are almost certainly this species. See the leaves of Trema micrantha (Cannabaceae), which are very similar but rough to the touch and usually with clearer teeth. Muntingia calabura (Muntingiaceae) also is similar. Three other genera of trees in the Grewioideae are found in Panama. Clappertonia has one species known only in w. Panama and seldom seen. Goethalsia is widely known in Costa Rica but very rare in Panama. The last genus, Triumfetta, has four species, but they are weak shrubs that only marginally can be called treelets. The leaves of Triumfetta resemble those of Heliocarpus, but have denser pubescence.

Malvaceae—Grewioideae

Luehea speciosa

Trichospermum galeottii

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Malvaceae—Malvoideae Hampea appendiculata

Hampea appendiculata

Ma Lvo IDeae (subfamily within Malvaceae). All genera that were in the old Malvaceae

are now in the subfamily Malvoideae in the newly expanded Malvaceae. Most of them are herbs, shrubs, or small trees, and they include the ornamental Hibiscus as well as the cotton plant, Gossypium. Most have wide, often lobed leaves, with palmate venation at the base, and they all have large flowers resembling those of the well-known Hibiscus. There are 21 malvoids in Panama growing as trees or treelets, and we cover 3.

Hampea appendiculata (playo blanco, azote).

A small tree of forest edge. Trunk is straight, cylindrical. Leaves are simple, alternate, with three major

veins (or even five) arising together from the base. There are two tiny brown swellings just above the base of the leaf, barely visible. Juveniles have much larger, wider leaves. Flowers are showy, white, with many stamens. Fruit are dry capsules that open into three valves. Distribution: A species of edge and natural clearings in the forest, throughout Panama and Costa Rica, in all habitats but the driest. Recognition: Lacking heart-shaped leaves like most Malvaceae, this tree is perhaps confused with Melastomataceae, all of which have three (or more) major veins but opposite leaves. The three major veins in Hampea are more prominent than in other Malvaceae except for those with really wide, lobed leaves. Juvenile Hampea with wide leaves could be mistaken for Heliocarpus (Grewioideae) or Ochroma (Bombacoideae). Hampea is very fast growing. Panama has five other Hampea species, but none of them are seen often. H. micrantha can be found in mountains around Cerro Jefe, and has glossy green leaves with venation similar to that of H. appendiculata.

Wercklea cocleana

Wercklea cocleana

Wercklea cocleana (amapola, papo de mon-

taña). A small tree of montane forest. Leaves are large, wide, rounded, with several main veins arising

from the heart-shaped base. They are toothed and sometimes slightly lobed. There are bunched green stipules at the base of each petiole, like tiny leaves. Flowers are large, pink, Hibiscus-like. Fruits are dry, fuzzy capsules. Distribution: In lower montane forest of c. to w. Panama and s. Costa Rica, at forest edges, roadsides, and clearings. Recognition: This should only be confused with other big-leaved Malvaceae. See Heliocarpus americanus (Grewioideae), which has very similar leaves but small yellow flowers, or the balsa, Ochroma pyramidale (Bombacoideae).

Malvaceae—Malvoideae

Hampea appendiculata

Wercklea cocleana

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Malvaceae—Sterculioideae Wercklea woodsonii

Wercklea woodsonii

Wercklea woodsonii (amapola, papo de montaña). A medium-sized tree of montane forest. Very similar to W. cocleana, but with yellow flowers and twigs that produce a gluelike sap when broken. Distribution: In lower montane forest of w. Panama and c.

Costa Rica. Often a roadside tree. Recognition: See the previous species. Two other Wercklea species are found in montane forests of w. Panama, one of which reaches Costa Rica. A third species is known in Caribbean lowland forests, but is seldom seen. All have large, round to lobed leaves. The Malvoideae include seven more genera in the Panama checklist, mostly treelets or small trees and nearly all with broad, heart-shaped leaves. One is the cotton genus, Gossypium (two species); the others are Abutilon (two), Lopimia (one), Malvaviscus (one), Pavonia (one), Talipariti (one), and Thespesia (one).

Sterculia apetala

Sterculia apetala

Ste Rcu LIo IDeae (subfamily within Malvaceae). This subfamily includes many of the

genera formerly classified in the family Sterculiaceae; see the Byttnerioideae for the rest. There are two genera of trees in Panama, one little known and with a single species. The other, Sterculia, includes large forest trees, one of which is the country’s national tree, el árbol panamá. All told, then, Panama has six species of sterculioids, of which we cover one.

Sterculia apetala (panamá). A tall, magnificent

tree of the Pacific slope. The trunk is straight, cylindrical, with large buttresses. Bark is smooth and yellowish. Leaves have five lobes, are often folded, and are clustered densely toward the ends of branches. Juvenile leaves are much larger than canopy leaves. The five-parted purple and yellow flowers appear to have petals, but they are really sepals. Fruits are woody capsules with orange hairs on the inside.

Distribution: A tree of the Pacific slope, widely known in farmland and often appearing in towns, growing on its own though also planted. There are many big trees in Panama City parks. Also reaches the Caribbean near the Canal, and there are very tall trees in old growth at Barro Colorado I. Juveniles only appear in open areas or natural forest clearings. Recognition: Large juveniles might be mistaken for Cecropia or especially Pourouma (both Urticaceae). Leaves of Pourouma and Sterculia have essentially the same shape, but the venation and color of the underside are quite different. Otherwise, S. apetala might only be confused with other Malvaceae with wide, lobed leaves; check Ochroma and Cavanillesia (Bombacoideae) or Heliocarpus (Grewioideae). Four more species of Sterculia are known in Panama, all from wet lowlands or lower montane sites; two reach Costa Rica. Two are quite common in c. Panama (Cerro Azul, Cerro Jefe, and Santa Rita). S. recordiana has wide, rounded leaves as an adult, but highly lobed leaves (like Pourouma) as a juvenile; they are bluish on the underside with three large veins arising from the base. The other genus in the Sterculioideae is Pterygota, with one species. It is known only in mountains of w. Panama and c. Costa Rica. It has wide, heart-shaped, but unlobed leaves.

Malvaceae—Sterculioideae

Wercklea woodsonii

Sterculia apetala

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Malvaceae—Tilioideae Mortoniodendron anisophyllum

Mortoniodendron anisophyllum

Tilioideae (subfamily within Malvaceae). This subfamily includes the remainder of the old

Tiliaceae, those that did not go into the Grewioideae. In North America and Europe, tilioids are known for the linden, or basswood tree, Tilia. In Panama, there is just one genus with two species, and we cover one.

Mortoniodendron anisophyllum. A mediumsized tree of wet forest. The trunk forks at low heights. Leaves are simple, alternate, narrow and pointed, but rounded at the base, where three main

veins emerge. The leaf is slightly asymmetric at the base (hence the scientific name). Flowers are white or yellow with five narrow petals. Fruits are spherical capsules that turn bright orange. d istribution: In wet forests of the Caribbean slope and the Osa Peninsula; sometimes in lower montane forests. Fairly common near the Canal and can be found easily at Fort San Lorenzo at the mouth of the Chagres River. Recognition: The narrow leaves with a pointed tip are quite different from those of most Malvaceae, and you might think of Euphorbiaceae at first. The asymmetric leaf base is a good character. Leaves are also similar to those of Gleospermum blakeanum, a small shrub in the Violaceae (not illustrated). Another species of Mortoniodendron is known from a few sites in Panama but is seen widely in wet lowlands of Costa Rica. It also has narrow leaves but lacks the asymmetric base, and will be easy to confuse.

Melastomataceae Bellucia pentamera

Bellucia pentamera

A very large tropical family of mostly treelets, shrubs, and herbs, with some tall trees and a few lianas. There are 4500 species worldwide and 3000 in the American tropics. It is the third largest family among trees of Panama, with 143 species (after Fabaceae and Rubiaceae), and it includes the largest genus, Miconia, with 65 species. We illustrate 12 Melastomataceae and describe 6 more. This is one of the easiest plant families to recognize. From the first Melastomataceae you look at you will nearly always know it. Leaves are always opposite, and there are (nearly) always three primary veins arising near the base that run parallel for most of the length of the leaf. In most species, there are additional parallel veins, often one more pair, that are slightly smaller than the three main veins. Between these five parallel veins and at right angles to them are a series of small tertiary veins packed in ladderlike formation. A few other species have three major veins running parallel for most of the leaf’s length, but none have five large parallel veins and none have such prominent, ladderlike tertiary veins. This dis-

tinctive leaf form, however, does not hold in one small group of Melastomataceae that must be learned separately (see Mouriri). Flowers of the family are distinctive, usually showy and fairly large, with stamens folded at an elbow. Recognizing species within the Melastomataceae, however, can be difficult because of the very large number of species. Moreover, genera are mostly distinguished by floral details and not easy to learn. Nevertheless, there are a large number of species with conspicuous features that you will have no trouble getting to know.

Bellucia pentamera (coronillo). A small forest tree. Leaves are large, thick, rounded, almost as wide as they are long. The leaf underside is a lighter than above, but lacks hairs or color. Branchlets and leaf petioles are grooved. Flowers are large, white or purple, produced along the branches. Fruits are large green berries. d istribution: Primarily along the Pacific coast, in moist or lower montane but not dry areas, and throughout the Canal Area. Nowhere especially common. Recognition: Another species with large, rounded leaves is Miconia argentea, but it has white leaf undersides. Some other species have fairly broad, green leaves, but not as wide as those of Bellucia. See Miconia hondurensis and Henriettea succosa, which have similar leaves, plus another species not illustrated, Henriettella fascicularis. A second Bellucia species also has large, thick leaves. It is seldom collected, but may be confused with B. pentamera.

Melastomataceae

Mortoniodendron anisophyllum

Bellucia pentamera

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Melastomataceae Clidemia octona

Clidemia octona

Clidemia octona (canillo). A small treelet of open areas. Only about 3 m tall. Stem, branches, and leaves are covered with a dense coat of fine, silvery hairs. Leaves are broad, heart-shaped, usually with seven main veins. Flowers are white and fruits are

blue berries. d istribution: Widespread in moist and wet lowlands plus occasionally at lower montane sites. Grows in disturbed areas and secondary forest. Recognition: Dense hairs and heart-shaped leaves are good characters. Another Clidemia, C. crenulata (not illustrated), is similar but has swollen leaf chambers that ants inhabit. In addition to those two, seven more species of Clidemia are found in Panama, and several of those are widespread forest or edge treelets throughout Panama and Costa Rica. Most Clidemia have a lot of hairs, often stiff hairs, and most have flowers on branches among leaves, not at the branch ends as in Miconia.

Conostegia bracteata

Conostegia bracteata

Conostegia bracteata (canillo). Not illustrated. A small tree of moist to montane forest. Leaves are pointed at both ends and have five major veins originating near the leaf base. There are long white hairs on veins below, and much shorter hairs on the lower leaf surface. Flowers are white, in terminal spikes; fruits are purple berries. d istribution: Widespread in moist to montane forests; rare in old forest at Barro Colorado I. and Soberania. Recognition: Conostegia is similar to Miconia in having terminal flower clusters, but is usually (but not always) distinguished by having hairs, as in Clidemia.

Conostegia cinnamomea

Conostegia cinnamomea

Conostegia cinnamomea (canillo). A treelet of wet to moist forests. Branches are low, shrublike.

Leaves are smooth, dark green above but light below. The inner pair of major side veins arises well above the leaf base, but the outer pair (right at the leaf margin) arises at the base. Flowers are small, white to pink, and fruits are dark berries. d istribution: In most habitats in Panama except the driest, and there are a few records in s. Costa Rica. Often along streams or slopes near streams. Recognition: Similar to Miconia nervosa, but the latter has seven main leaf veins. M. affinis and M. centronioides have similarly shaped leaves, but all their main veins arise at the leaf base.

Melastomataceae

Clidemia octona

Conostegia cinnamomea

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Melastomataceae Conostegia rufescens

Conostegia rufescens

Conostegia rufescens (canillo). Not illustrated. A medium-sized cloud-forest tree. Trunk is straight, cy-

lindrical, like a real tree (unlike other Conostegia), with smooth, white bark. There are sparse red hairs on twigs. Leaves have five main parallel veins, the second pair arising above the leaf base, as in C. cinnamomea. Veins underneath are reddish but without hairs. Flowers are white, with yellow stamens, at the ends of branches. d istribution: Principally a lower montane species, throughout Panama and Costa Rica, but also found in wet lowlands. Recognition: Much larger than most melastomes, and when a tree, the striking white bark is very distinctive.

Conostegia speciosa

Conostegia speciosa

Conostegia speciosa (canillo). A weedy treelet.

Leaves have seven main veins arising well above the leaf base, and fine white pubescence. Flowers are purple, with yellow stamens. d istribution: Known widely but sparsely in open areas of lowlands; very common around Panama City. Recognition: Like C. bracteata, but veins arise at leaf base in that species. Soft white hairs are not found in many Miconia.

Conostegia xalapensis

Conostegia xalapensis

Conostegia xalapensis (papelillo, dos caras, raspa lengua, quita manteca). A small tree of wet and cloud forest. Bark is light colored and flakes off in small pieces. Leaves are narrow, pointed at both ends, toothed, and a rich golden to whitish color beneath (from a tiny coat of hairs). Flowers are in termi-

nal clusters, and are white with yellow stamens. d istribution: An edge species in lower montane forest and in the wet Caribbean lowlands. Recognition: Miconia elata is quite similar, but the leaf is larger and lacks teeth. See also M. argentea. Sixteen more species of Conostegia are known as treelets in Panama. Moreover, the distinction between Conostegia and Miconia is a minute floral trait, so vegetatively, you can think of them as one group, though Conostegia tends to be hairy and Miconia not (but there are plenty of exceptions). There are lower montane species, a few in the lowlands, some are quite common and widespread, but several are seldom seen and poorly known.

Melastomataceae

Conostegia speciosa

Conostegia xalapensis

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Melastomataceae Henriettea succosa

Henriettea succosa

Henriettea succosa (canillo). A small tree of for-

est edge. Leaves are broad but pointed at both ends and light brownish gold below. Upper leaf surface is rough to the touch, and the lower surface and margin are densely pubescent. Flowers and fruits are borne on the branches, below or among leaves. d istribution: Mostly known in the Canal Area, where it can be found widely, including Pipeline Rd. Otherwise known at a few wet or lower montane sites. Usually a forest-edge species. Recognition: The rough upper leaf surface is a good clue. A similar

species, not illustrated, is Henriettella fascicularis, a slightly larger tree with similar leaves; H. fascicularis lacks hairs on its leaf margin, though. This species illustrates a key difference among genera of Melastomataceae: in Henriettea, most Clidemia, and several other genera, flowers are borne on branches below leaves, whereas in Miconia and Conostegia (the two huge genera), flowers are always in clusters at the ends of branches. This often helps, because even without flowers and fruits, Henriettea will show scars from fallen flowers on the branches. The genus contains one more species, Henriettea strigosa. It is known only on mountains near Panama City (perhaps just one record). As a seeming effort to maximize confusion, there is a separate genus Henriettella, which in Spanish takes exactly the same pronunciation. It is similar in having flowers on branches; there are five species known in Panama, and all but one are very rare.

Miconia affinis

Miconia affinis

Genus Miconia. This is the largest genus of trees in Panama (65 species) and the largest in the American tropics (1000 species). Moreover, Panama has 35 more species too small to be counted as treelets (though this distinction is imprecise) and even some vines. The 65 we include in the checklist are treelets or small trees, with one just barely a canopy-sized tree. All Miconia have white flowers in terminal clusters, and many flower after rains during the dry season. All produce small berries that attract many birds. Most grow at the forest edge or natural clearings inside mature forest. Many are montane. We include photos of six species and mention three

others for comparison, which unfortunately leaves a very large number of similar small trees uncovered. Moreover, the genus Conostegia (21 more treelets) is essentially the same as Miconia, differing only in a floral detail. Here we come close to covering most Miconia that you will encounter in lowland sites in easy-to-visit places, but you will still have to be cautious about naming this group to species; in montane sites or wet Caribbean coast, you are very likely to encounter species we do not cover here. Indeed, an entire book could be devoted to this genus alone.

Miconia affinis (canillo). A small tree of forest and edge. Leaves are fairly large, narrow and pointed at both ends, and lack hairs of any sort; the five parallel veins all arise near the base of the leaf. d istribution: Found in wet and moist zones, also lower montane areas, very common near the Canal. As Miconia go, this species tolerates shade in mature forest, but it favors clearings and can be seen at forest edge and along roads. Recognition: Similar to Conostegia cinnamomea, but that species has two main veins arising well above the leaf base.

Melastomataceae

Henriettea succosa

Miconia affinis

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Melastomataceae Miconia argentea

Miconia argentea

Miconia argentea (papelillo, dos caras). A medium-sized tree of forest edge and clearings. This is large as melastomes go, just barely reaching canopy size in mature forest. Leaves are large, almost round, and have conspicuous white undersides. In bigger individuals, the whitish brown bark peels away in vertical, papery strips. Flowers are small, white, and carried in dense, pyramidal clusters above the leaves. The flowering pattern of this species is remarkable.

For several weeks, trees gradually develop flower buds, and clusters of dull gray buds are visible throughout the dry season. Then on one day in January or February, many individuals over large areas complete flower development, and bright white flower clusters appear on nearly every tree. The following day, the flowers start dropping, and the dull gray color of the clusters is all that remains by the second day. Berries are small and dark blue. d istribution: An abundant and familiar tree of roadsides, farmlands, and towns, especially on the Pacific slope. Within the forest, it only occurs in natural clearings where there is light, but it is still fairly numerous. One of the most abundant species of secondary forests in the Canal Area. Recognition: Quite possibly the easiest tree in Panama to learn. Another species that might confound is M. elata (not illustrated), but its leaves are narrower and the color underneath is golden, not white.

Miconia centronioides

Miconia centronioides

Miconia centronioides (papelillo, dos caras, canillo). A small cloud-forest tree. Leaves are fairly

large, pointed at both ends, and a dark brown color underneath (from small hairs). d istribution: Only found in mountains of c. and e. Panama, but can be common. Mainly an edge and secondary species. Recognition: Similar to M. elata (not illustrated), which is mainly a lowland species, but its leaves have a lighter, more golden color. Two other species, M. dodecandra and M. serrulata (not illustrated), have similar leaf color, but have round or heart-shaped leaf bases; M. dodecandra is montane, but M. serrulata also occurs in lowland wet forests.

Miconia elata

Miconia elata

Miconia elata (papelillo, dos caras, canillo). Not illustrated. A small tree of forest edge. Leaves are

broad, sometimes almost round, but the base comes to a point (though perhaps blunt). They are light brown below, with five main veins arising right at the leaf base. d istribution: Mostly known near the Canal, but widespread from Pacific to Caribbean and lower montane regions in this area. Has also been found at a few sites in w. Panama and Costa Rica. Recognition: Similar to M. argentea, but leaves are brownish below rather than white; the difference can be subtle. M. serrulata (not illustrated) is a similar species, but the leaf base is heart-shaped. See also M. centronioides.

Melastomataceae

Miconia argentea

Miconia centronioides

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Melastomataceae Miconia hondurensis

Miconia hondurensis

Miconia hondurensis (canillo). A small forest

Miconia impetiolaris

Miconia impetiolaris (dos caras, oreja de

tree. Leaves are fairly large, pointed at both ends, smooth shiny green, darker above than below. d istribution: Widely distributed but seldom seen. Recognition: Similar to M. affinis, but leaves are much shinier.

Miconia impetiolaris

mula). A treelet of forest edge. Oval leaves have cream- or copper-colored undersides; the upper side of the leaf is shiny green. The base of each leaf extends back along the petiole, and folds against the stem. It flowers in January or February, following the pattern of Miconia argentea; indeed the two species sometimes flower on the same day. d istribution: Widely known, but most common in the Canal Area, where it is a widespread forestedge and roadside species. Recognition: The folded leaf base separates this from other melastomes.

Melastomataceae

Miconia hondurensis

Miconia impetiolaris

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Melastomataceae Miconia nervosa

Miconia nervosa (canillo). A small forest tree. Leaves are pointed at both ends, light blue-green below. There are seven parallel veins: one small pair at the margins that arise at the leaf base, and two more pairs arising well above the leaf base. d istribution: In wet lowlands throughout Panama and Costa Rica. This species handles shade a bit better than other Miconia and is seen in the forest, though still favors gaps and clearings. Recognition: Similar to Conostegia cinnamomea, but has an extra a pair of leaf veins. See also M. affinis and M. hondurensis.

Miconia nervosa

Miconia oinochrophylla

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let with red leaves. Has typical melastome venation, with narrow, pointed leaves whose upper side is dark green. The underside, however, is a remarkable deep purplish red. d istribution: In wet and lower montane forests of c. Panama and the Osa Peninsula, plus a few other locations. Recognition: The leaf color is unmistakable.

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Miconia trinervia

Miconia trinervia

Miconia trinervia (canillo). A small tree of montane forest. Leaves are shiny green above, thick, and only have three main parallel veins. d istribution: Sometimes lower montane, and common around Cerro Jefe near the Canal, but also found in very wet lowland forests. Recognition: Other melastomes we have described have five or more veins running parallel for the length of the leaf.

Melastomataceae

Miconia nervosa

Miconia trinervia

309

310

Meliaceae

Mouriri myrtilloides

Mouriri myrtilloides

Mouriri myrtilloides (guayabillo, solacra, cierito,

sequara). An understory treelet not at all resembling

other Melastomataceae. The small stem is straight and vertical, and pairs of horizontal branches and branchlets emerge opposite one another. Leaves are opposite, regularly spaced and in a flat plane, with many per branch; the branches are long. Leaf margins are undulate, and the leaves are small, narrow, and pointed. Secondary veins are inconspicuous. There is no petiole, so the leaves grow right against the branch. Branchlets are slightly winged, or at least with small ridges. Flowers are white with yellow stamens and are produced along branches between leaf pairs or branchlet pairs, and fruits are dark berries. d istribution: Throughout Panama and Costa Rica in moist and wet zones, sometimes in lower mountains. Abundant in old growth at Barro Colorado I. Strictly a species of the forest interior. Recognition: Venation is utterly different than in its relatives, so this species will not be learned with Melastomataceae. Nevertheless, it is easy to recognize, at least when you look

closely and see that there is no petiole. It can be confused with Eugenia nesiotica (Myrtaceae), which also has opposite leaves with an undulate margin, but Eugenia and all the Myrtaceae have petioles, although sometimes rather short ones. Moreover, most Myrtaceae have tiny translucent spots in the leaves, which Mouriri lacks. See also Lafoensia (Lythraceae) plus Vochysia and especially Qualea (Vochysiaceae). Five other Mouriri species are found in Panama. M. panamensis is only found in wet forests east of the Canal and is a canopy tree; three others are sparse and better known in Costa Rica, and a fourth is restricted to far e. Panama. They all have smooth, nearly veinless leaves on very short petioles, but some can be big trees, and they generally lack the wavy margins so typical of M. myrtilloides. There are still 13 genera of Melastomataceae we have not covered: Adelobotrys (one species), Axinaea (one), Blakea (six), Graffenrieda (four), Leandra (three), Loreya (one), Meriania (five), Ossaea (three), Tessmannianthus (three), Tibouchina (one), Tococa (one), Topobea (one), and Votomita (one). The latter is in the Mouriri group, and has narrow, smallish leaves with inconspicuous venation. All the rest have prominent venation like all the other melastomes, and for the most part, these genera do not have easy features to separate them from the ones we did illustrate. Several are largely epiphytic but sometimes grow as trees.

Carapa guianensis

Meliaceae

Carapa guianensis

A moderate-sized tropical family most of which are trees, important both ecologically and economically. It is the family of mahogany, and “true” mahogany is the American genus Swietenia (there are also African mahoganies in different genera), and several other species in the family produce valuable timber. Meliaceae has 550 species worldwide and 130 in the American tropics. Panama has 31 of those, and we cover 11. Meliaceae has a consistent set of leaf characters that make it fairly easy to learn. The entire family has compound leaves, and the petiole has a brown swelling at the base. Unlike the Fabaceae, the swelling is not cylindrical, but is flattened or even indented above. Burseraceae, Anacardiaceae, and Sapindaceae all have the same kind of petiole. Most

Meliaceae are even-pinnate, with leaflets in opposite pairs and no terminal leaflet; however, the major genus Trichilia has odd-pinnate leaves, sometimes with alternate leaflets. Flowers and fruits of Meliaceae are distinct only in detail and mostly not useful for identification, with the exception of the winged seeds of Swietenia and Cedrela.

Carapa guianensis (tangaré, cedro bateo). A huge canopy tree of wet areas. The trunk is straight, cylindrical, with substantial buttresses in big trees. The bark is brown and peels off in small circles. Leaves are even-compound and alternate, clustered toward the ends of branches. Leaflets are large, opposite, rounded at the tip and with a small notch, usually six to eight per leaf. Juvenile stems are unbranched and have large leaves emerging helically from the stem. Flowers are small, white or yellowish. Fruits are large, round woody capsules, with four valves that split open to reveal pulp around the seeds. d istribution: Lowland wet forests of the Caribbean slope and the Osa Peninsula, plus a few lower montane sites. It is most often seen in swampy areas or along rivers near the Caribbean coast. Not seen along roads or in open areas. Recognition: Leaves lacking a terminal leaflet, and with a petiole

Meliaceae

311

Mouriri myrtilloides

Carapa guianensis

that is swollen, brown, and flattened, eliminate Trichilia and most other compound-leaved families. Carapa has fewer, larger, and rounder leaflets than Cedrela, Guarea, or Swietenia; but see Guarea grandifolia for comparison. Carapa is harvested commercially for timber.

Recent research has detected an additional Carapa species known now at just two locations on the Caribbean slope, one in far n. Costa Rica and one in e. Panama.

312

Meliaceae

Cedrela odorata

Cedrela odorata

Cedrela odorata (Spanish cedar, cedro, cedro

amargo). A tall timber tree. Large trees have plank buttresses at the base. The bark is light brown or gray, with vertical fissures. Leaves are even-compound with 10–22 opposite leaflets that have tiny pits (domatia) on the underside at the base of the secondary veins. All young parts of the plant are fuzzy and smell strongly when crushed. Big trees are deciduous for part of the dry season. Flowers are small and white. Fruits mature for nearly a year, then the woody capsules split open into a star shape. Inside are seeds with large wings. d istribution: Primarily on the Pacific side of the isthmus throughout, but nowhere common. It can be found regularly in secondary for-

ests around Panama City, and sometimes along roads or in towns. Most trees are less than 50 cm in diameter with only small buttresses. Recognition: Very similar to Swietenia macrophylla, and the two are often confused; note that Cedrela usually has more leaflets that are symmetric (or nearly so) and rounded at the base. Both species have fissured bark, but are quite different in appearance. The cedro has fine timber, is usually considered next in value in the New World after mahogany. It may have once been more common on the Pacific slope of Panama; large individuals probably would have been harvested. Currently not common and there is no commercial harvest. The cedro can grow rapidly when it has light, but small saplings are frequently attacked by a moth larva when they are cultivated, and this has prevented production in plantations. Two more species of Cedrela are found in Panama. C. fissilis is quite common, usually at higher elevations, and is similar to C. odorata, but with larger leaves. The other species is known only in mountains of w. Panama and Costa Rica.

Guarea grandifolia

Guarea grandifolia

Guarea grandifolia (chuchupate, cedro macho,

tres bocas). A large forest tree. The trunk is straight, buttresses at best small. Leaves are even-pinnate, typically with 10–12 opposite leaflets with pointed

tips. There is a small bud at the end of the leaf that extends beyond the final pair. This bud is capable of growth, and new leaflets can be added to the leaf. Brown fruits are capsules shaped like small pears that split open to reveal seeds. d istribution: Common throughout the Canal Area; otherwise known from a few widely scattered locations. Only inside the forest. Recognition: Guarea is the major genus with even-pinnate leaves. The growing tip at the end of the leaves is distinctive. Look carefully, because it is small, and it may be missing from some leaves. G. grandifolia has somewhat larger and fewer leaflets compared to its congeners, but see G. guidonia.

Meliaceae

Cedrela odorata

Guarea grandifolia

313

314

Meliaceae

Guarea guidonia

Guarea guidonia

Guarea guidonia (chuchupate, cedro macho). A medium-sized forest tree. The trunk is straight but branches often. Leaves are even-pinnate, typically with 8–10 opposite leaflets that are narrow and have

pointed tips. There is a small bud at the end of the leaf. Leaves on small saplings can be much larger, with many more leaflets. The fruit capsule has four or five valves, and seeds within are covered with an orange-red pulp. d istribution: Common throughout the Canal Area and to the east. Two other records as far as n. Costa Rica suggest it is widely distributed. Only inside the forest. Recognition: Look for the bud at the end of leaves. G. guidonia has many smallish leaflets. Talisia (Sapindaceae) has similar leaves and might be mistaken for Guarea. See also Inga (Fabaceae—Mimosoideae), but all Inga have glands on the rachis.

Guarea pterorhachis

Guarea pterorhachis

Guarea pterorhachis. A medium-sized wet-forest tree. The trunk is straight and buttresses at best are small. Bark peels off in small pieces. Leaves are even-pinnate, and the rachis is winged. There is a small bud at the end of the leaf. The brown fruits have two valves and a red berry within each.

d istribution: From wet lowlands and occasionally lower montane sites. Fairly common at Santa Rita east of the Canal. Recognition: Not confused with other Guarea because of the winged rachis. At a glance looks like the many Inga (Fabaceae—Mimosoideae), which have wings on the rachis, but all Inga have glands at the base of each leaflet pair. Guarea has six more species in Panama, all sharing even-pinnate leaves with a growing bud at the tip. G. bullata and G. glabra are common around the Canal and in lowland forest in general; they resemble G. guidonia but with fewer leaflets. Other species are widespread in wet and lower montane forests, though one is very rare.

Meliaceae

Guarea guidonia

Guarea pterorhachis

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Meliaceae

Swietenia macrophylla

Swietenia macrophylla

Swietenia macrophylla (caoba, mahogany). A

magnificent tall tree mostly of parks and cities now. Buttresses are huge in big trees. Bark is gray, with vertical fissures. The crown can be immense, wide, round, tall, with leaves from near the ground to very high. Leaves are even-pinnate, bunched toward the ends of branches, and leaflets are often not quite opposite one another. Each leaflet is asymmetric, much narrower on the trailing edge. Flowers are tiny, white to greenish, in clusters above the leaves. Fruits are brown capsules held erect over the leaves, easily visible before they open. The valves then crack off and large, winged seeds float off like helicopters.

d istribution: Now mostly a street tree, planted in cities and towns everywhere. There are still said to be wild populations in remote forests of e. Panama, far from roads. We know a handful of trees in secondary forest not far from Panama City, but these may be descended from ornamentals. Where it does occur naturally, juveniles are scarce and always in open areas, but adults can be immense canopy trees inside the forest. It is a species of drier areas elsewhere. Recognition: Easy to learn in cities from the buttresses and bark. Leaves are very similar to those of Cedrela, and juveniles of both species can be found in towns or farms, but only Sweitenia is planted along streets. Leaflets of Swietenia are more pointed at the base and very asymmetric compared to those of Cedrela. If you do come across a mahogany in the forest, it will most likely be a large one, and buttresses and bark would be the features cluing you in. The caoba was presumably more common around Panama City and on the Pacific slope before it was harvested. Spanish reports from 300–500 years ago describe huge wooden structures made from it.

Trichilia hirta

Trichilia hirta

Trichilia hirta (conejo colorado, mata piojo). A medium-sized tree of the dry zone. Bark is brown or reddish. Leaves are very long, with many pointed leaflets; they are not quite opposite along the rachis (subopposite). There is a terminal leaflet, and it usually is angled slightly from the rachis; sometimes this effect makes a leaf look even-pinnate, because the final leaflet is midway between being terminal and be-

ing paired with the next leaflet. Twigs and leaf undersurfaces have short and inconspicuous pubescence. Fruits are woody capsules, with three valves that open to reveal seeds carrying bright red arils. d istribution: In the driest Pacific zone of c. Panama and nw. Costa Rica, where it appears in secondary forest and agricultural areas. Quite common on the Pacific side of the Canal Area. Recognition: The long leaves with many leaflets should recall Spondias (Anacardiaceae), which is abundant in agricultural areas, but Spondias lacks pubescence and leaflets are more precisely opposite. Mosquitoxylon is another anacard that does have pubescence, but it is a rare, wet-forest tree. The terminal leaflet angled slightly to one side is a good Trichilia trait, though not universal in the genus. See T. martiana, which has similar but more-pubescent leaves.

Meliaceae

Swietenia macrophylla

Trichilia hirta

317

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Meliaceae

Trichilia martiana

Trichilia martiana

Trichilia martiana (conejo colorado, mata piojo). A medium-sized tree of dry forests. The trunk is slightly ridged. Leaflets can be precisely opposite, or subopposite, about seven to nine per leaf. The end leaflet is usually slightly angled to one side, and sometimes appears paired with another leaflet. Secondary veins are conspicuous below, yellowish, and

tightly parallel. d istribution: Known principally in riverside forests of drier areas of the Pacific coast, and on limestone near the Canal (Madden Dam), but it is widely collected along the Pacific slope and even lower mountains in Costa Rica. Recognition: As in many Trichilia, the end leaflet is slightly angled. Among the Trichilia, T. martiana is most like T. pleeana, which is also found in drier areas but has fewer leaflets. T. hirta likewise grows in dry habitats, but has small and narrow leaflets. Leaflets of T. martiana resemble those of Lonchocarpus heptaphyllus (Fabaceae—Papilionoideae), but Lonchocarpus always has strictly opposite leaflets (which T. martiana is inconsistent about). There are other Fabaceae that might be confused, but remember that Fabaceae have cylindrical petiole bases. See also Spondias and Tapirira (Anacardiaceae).

Trichilia pallida

Trichilia pallida

Trichilia pallida (terciopelo, conejito colorado). A medium-sized forest tree. The trunk is straight, some-

what irregular. Bark is cream-colored or yellow. Leaves have three to seven leaflets usually opposite (or subopposite) along the rachis, with the terminal leaflet larger than the others and sometimes angled to the side. On occasion, a leaf with only the end leaflet appears, becoming for all appearances a simple leaf. d istribution: Found widely in moist lowland to lower montane forest, never common. Recognition: The enlarged end leaflet does not look like those of other Trichilia. Some Fabaceae have this appearance (see Lonchocarpus), but with a very different petiole base.

Meliaceae

Trichilia martiana

Trichilia pallida

319

320

Meliaceae

Trichilia pleeana

Trichilia pleeana

Trichilia pleeana (alfajía, fosforito, alfaje). A medium-sized tree of drier regions. Bark is white. Leaves

are odd-pinnate, with five to seven leaflets that can be alternate or subopposite along the rachis. The terminal leaflet often angles to one side, almost becoming a side leaflet. d istribution: Collected widely but not well known. Grows on the Pacific side of the Canal Area, is common on limestone formations (Madden Dam and Fort Sherman), and appears in dry nw. Costa Rica. Recognition: When the end leaflet is angled, it clearly is a Trichilia, and somewhat like T. tuberculata but with wider leaflets. Might recall Sapindaceae or Fabaceae.

Trichilia tuberculata

Trichilia tuberculata

Trichilia tuberculata (alfajía, alfajía colorado, fos-

forito, alfaje). A canopy tree of mature forest. Large trees have straight, cylindrical trunks, with no buttresses, and are branched only near the top. The bark has small, white “tubercles” (actually lenticels to be precise), each consisting of a pair of vertical white bars with a black line in between. Leaves are oddcompound, and leaflets are narrow, dark shiny green, and mostly alternate along the rachis. Outer leaflets are larger than inner ones. The end leaflet always angles to the side and sometimes appears to be paired with the next leaflet. There are often small, black circles on leaves, but this is a fungus and is not always present. Fruits are capsules that open when mature to reveal small, dark seeds with a bright red aril. d istribution: Abundant in the old forest of Barro Colorado I., where it is one of the dominant trees of

the canopy and is very common as a sapling. Found throughout the Canal Area, but less common. Elsewhere it is not so well known, though collected widely. Never seen outside the forest. Recognition: In large trees, the white lenticels are good characters but will only be noticed with close inspection. The angled end leaflet is obvious in saplings, but other Trichilia have this. Compared to others in the genus, T. tuberculata has narrow leaflets. This species, as well as some others of the genus, are called fosforito because the wood burns well when still wet (fosforito means little match). Six more Trichilia species are known in Panama. Several are widespread and fairly common, two from dry forest, one lower montane, and others from wet forest. You should be able to recognize the genus, given alternate to subopposite leaflets that are obviously not Fabaceae. One exception, however, T. trifolia, has both simple and trifoliate leaves, the latter like those of Allophylus (Sapindaceae). One other genus of Meliaceae, Ruagea, has two species known in mountains of w. Panama and c. Costa Rica. Leaves are like those of Trichilia, with an angled terminal leaflet, and a few Trichilia species occur in the montane habitat of Ruagea.

Meliaceae

Trichilia pleeana

Trichilia tuberculata

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Moraceae

Moraceae

Brosimum alicastrum

Brosimum alicastrum

not look like flowers. Fruits have the same globular shape.

Brosimum alicastrum (breadnut, berbá, cacique). A tall tree of moist-forest canopy. The trunk is

An important tropical tree family, with 1050 species worldwide and 270 in tropical America. The fig genus Ficus accounts for more than half of those species. Most are trees, but many Ficus are hemiepiphytes, or strangler figs, and there are a few lianas and herbs. Panama has 74 tree species, of which we illustrate 13 and describe 1 other. The Moraceae have consistent traits that are easy to learn. They have simple, alternate leaves, and these tend to be spaced evenly along branches and in a flat plane. Many Moraceae have dark green leaf undersides with contrasting and finely reticulate yellow veins, and leaves of the family have a general appearance that is not hard to learn. The best trait is the sap: nearly all species produce white latex at wounds, and many species gush copiously. Another useful trait is the stipule scar. In many species, the stipule surrounds the apex of the twig, and when it falls off it leaves behind a circular scar at the base of each leaf; some Moraceae lack this trait, however. Flowers are very peculiar, nearly always tiny, packed together in a globular head; they do not open and do

straight and cylindrical, and big trees have prominent buttresses with rounded tops. The bark is smooth and light gray. Branchlets have circular scars at each leaf base. Leaves are simple, alternate, and have yellow veins on the underside set against a dark bluish hue; even the smallest veins are yellow, creating a fine network against a dark background. There is a conspicuous collecting vein running along the leaf margin. In saplings, leaves are rough to the touch and can be toothed. Broken leaves gush white latex. Flowers are tiny, white, in globular clusters, and fruits are green, with single seeds. d istribution: Primarily a Pacific coast species, including fairly dry areas, but it is very common everywhere in the Canal Area. Not often seen along roads or in open areas. Recognition: Pseudolmedia spuria (not illustrated) and Sorocea affinis have leaves like those of B. alicastrum, more so than the other Brosimum, but they lack the strong contrast of light yellow veins against dark blue leaf underside and they are never large trees. The trunk and buttresses of B. alicastrum resemble those of Ficus insipida, the abundant fig, but the leaves are much different and can usually be found on the ground. The nuts of B. alicastrum are edible after boiling, and were once an important food to the Mayans in s. Mexico.

Brosimum guianense

Brosimum guianense

Brosimum guianense (berbá, cacique). A tall

wet-forest tree. The trunk can be huge, with large buttresses. Branchlets have circular scars where stip-

ules fell off. Leaves are simple, alternate, with a pointed tip, dark green above but lighter below with a silvery sheen. Veins are yellowish below, and there is a conspicuous collecting vein running along the leaf margin. Tertiary veins form a reticulate network of little squares. Broken leaves or twigs drip ample white latex. Flowers are yellow, on small, flat, mushroomshaped heads. d istribution: Lowland wet forests, sometimes in lower mountains. Found exclusively in mature forest, not open areas. Recognition: The leaves look solidly Moraceae, with the extended tip and reticulate veins, plus latex; the silvery sheen on the leaf underside distinguishes this species.

Moraceae

Brosimum alicastrum

Brosimum guianense

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Moraceae

Brosimum utile

Brosimum utile

Brosimum utile (sande, mastate, palo de vaca). A canopy tree of wet forests. The trunk is straight, cylindrical, with only small buttresses at the base. Bark is brown or reddish with white lenticels. Leafs are simple, alternate, lightish green on the underside with contrasting yellow veins. The stipules wrap around the growing end of each branchlet, but they soon fall off, leaving a scar ringing the branchlet at the base of

each leaf. Broken leaves or slashes in the bark produce drops of white latex. Flowers are tiny, on small white heads at the base of branches. Fruits are fleshy spheres, turning red, with white lenticles. d istribution: Lowland and lower montane wet forests, just reaching s. Costa Rica. Not a species of edge or open areas. It is the dominant canopy tree near Fort Sherman. Recognition: The leaves are longer and lighter in color than those of other Brosimum species such as the widespread B. alicastrum. Instead, B. utile may be confused with another Moraceae, Maquira guianensis, which also has long and light-colored leaves. But Maquira is a small tree with gray, not brown, bark. Brosimum has three more species in Panama, all with similar wet-forest distributions. They have stipule scars and lots of latex.

Castilla elastica

Castilla elastica

Castilla elastica (caucho, mastate blanco, hule).

Not illustrated. A roadside tree around Panama City. The trunk is straight and vertical, and branches are horizontal. Leaves are simple, alternate, spaced regularly along branches and in a flat plane; sometimes they have teeth. Leaves and twigs have dense, fine pubescence. During the dry season, there are usually some yellow leaves, and many trees are completely

deciduous. White latex flows from any cut or break, and scars ring the branches at each leaf base. Flowers are tiny, in small dense heads that look like fungus. d istribution: Abundant along roads near Panama City. Otherwise widely but sparsely known. Recognition: Leaves are similar in shape, size, and pubescence to those of Perebea xanthochyma, but leaves of Perebea narrow to a point at the base, whereas leaves of Castilla are broad at the base; Perebea also has much more prominent teeth. Castilla resembles two other common roadside species, Annona spraguei (Annonaceae) and Apeiba tibourbou (Malvaceae—Grewioideae), but neither of those has latex. Castilla was used as a source of rubber by Native Americans. A second Castilla species is known at only a few sites in Costa Rica and Panama.

Ficus insipida

Ficus insipida

Ficus insipida (fig, higuerón). A large, buttressed tree of roadsides. The bark is smooth and graybrown, and the buttresses have rounded tops. Leaves are bright, shiny green, with yellow veins, and secondary veins are closely spaced and tightly parallel. Fallen leaves turn bright yellow on the ground. The tip of each branchlet has a long, yellow, pointed stipule, which falls as the branch grows past it; this leaves a circular scar on the branch at the base of the

leaf, and the long yellow stipules are easy to find on the ground beneath big trees. Broken leaves drip white latex rapidly. All figs have flowers and seeds inside small, green balls. They never open, and tiny, specialized wasps pierce the balls to insert their own eggs (and in the process, pollinate the flower). d istribution: Common and conspicuous in open areas in all zones: along roads, farms, forest edge, and rivers. It is one of the characteristic trees of secondary forest, and often the largest tree in those habitats. It occasionally appears as a large tree in old-growth forest, but does not reproduce inside the forest; juveniles are seen only in open areas. Recognition: One of the first trees to be learned anywhere in Latin America. On big trees, the combination of large, smooth buttresses and bright green leaves with yellow veins make it easy to recognize. Check for fallen yellow leaves and the long yellow stipules on the ground to confirm identification in the forest. As juve-

Moraceae

325

Brosimum utile

Ficus insipida

niles, circular scars at the base of each leaf, copious white latex, and pointed stipules pin down the fig genus (Ficus); the large, bright, shiny leaves of F. insipida are key, but there is a very similar species of the Caribbean slope, Ficus yoponensis, which we do not illustrate. F. yoponensis has smaller leaves and stip-

ules and does not occur abundantly in secondary forest the way F. insipida does. Brosimum alicastrum has a trunk just like that of F. insipida, but has different leaves. See also Sapium glandulosum (Euphorbiaceae), which has figlike leaves and latex but very different bark and a conspicuous pair of leaf glands.

326

Moraceae

Ficus tonduzii

Ficus tonduzii

Ficus tonduzii (fig, higuerón). A large forest tree. Trunk and bark are similar to those of F. insipida, with all the fig traits (latex, circular branch scars, and yellow veins). Leaves are dull green and there are few, widely spaced secondary veins. d istribution: Widespread in lowland wet forest, less so in lower mountains. Often in valleys or along streams. It requires clearings within the forest, so juveniles are not seen in the shade, but it is not a secondary forest and edge species like F. insipida. Recognition: Trunk resembles that of the other big figs (F. insipida and F. yoponensis), but leaves are larger, duller, with much different venation. See

leaves of Poulsenia armata, very similar but with spines along the vein and on the petiole, and also Maquira guianensis. Ficus is one of the biggest genera of trees in Panama, with 39 native species. Many of these are stranglers, but are counted as trees because when their host dies they stand on their own trunks. The stranglers are difficult to identify because as saplings they are in the canopy, and as big trees their leaves are nowhere near the ground. Although quite variable in leaf form, they all have pointed, cone-shaped stipules at the ends of new branches, circular branch scars, and lots of white latex. You can get lots of practice with your fig identification because a number of species are planted in gardens, towns, and as houseplants. In cities of Panama, you can find F. benjamina, a species with small leaves, often grown in houses and gardens; F. elastica, the Indian rubber, which has broad, smooth leaves; and F. benghalensis, the banyan tree, with multiple stringy roots hanging from branches. All of those species are native to Asia.

Maclura tinctoria

Maclura tinctoria

Maclura tinctoria (moro, mora, amarillo). A large dry-forest tree. The trunk is straight, cylindrical, and large individuals have conspicuous swellings at the base extending onto roots that run along the ground. The bark is yellowish, with irregular, cream-colored

horizontal bands and lenticels, and small, lumpy spines. The leaves are simple, alternate, conspicuously toothed, asymmetric at the base, with a long pointed tip. White or cream-colored latex drips rapidly from broken leaves or bark slash. Male flowers are tiny, white, densely arranged on a long, narrow spike. Female flowers and fruits are in a globular heads. d istribution: Pacific coast of Panama and Costa Rica, most often seen in the driest zone, including pastureland. Also on limestone outcrops at Madden Dam near the Canal, where it is common. Recognition: On bigger trees, the trunk is conspicuous. Leaves are also distinctive, because teeth and latex seldom go together.

Moraceae

Ficus tonduzii

Maclura tinctoria

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Moraceae

Maquira guianensis

Maquira guianensis

Maquira guianensis (palo de pico). A medium-

sized tree of wetter areas. The bark is gray and twigs brown with white lenticels. Leaves are simple, alternate, in a flat plane, with a slightly asymmetric base and an extended narrow tip. Veins are yellow below,

and secondaries are few and widely spaced; adjacent secondary veins have a curving connector between them near the leaf margin. Any broken part produces lots of white latex. Ring scars at the base of leaves are weak. Flowers and fruits are in small, funguslike heads. d istribution: In moist, wet, and lower montane areas throughout. Recognition: Lacks encircling twig scars and the dark leaf underside of many Moraceae, but the white latex, yellow veins, and long tips are good family characters. Leaves are very similar to those of Ficus tonduzii, with widely spaced secondary veins, but note the extended tip in Maquira. One of the Moraceae we do not illustrate, Clarisia biflora, has fairly similar leaves.

Naucleopsis naga

Naucleopsis naga

Naucleopsis naga (palo de pico). A medium-

sized wet-forest tree. At the growing end of twigs, there is a short, pointed stipule, and at the base of each leaf there is a prominent scar encircling the

twig (where the stipule fell). Leaves have yellow veins below and a weakly pointed tip. Any broken part produces lots of white latex. Flowers are tiny, in dense balls; fruits are spheres covered in red spines. d istribution: In lower montane and lowland Caribbean forests, but not common. Recognition: Looks just like a Ficus, with pointed stipules, encircling scars, and latex. Differs in longish leaves and the fruit. Three other species of Naucleopsis are known in Panama. All are sparsely distributed in wet or montane forest throughout the region. They share the short, encircling stipule and latex; N. ulei is an easy one to learn, having very long, narrow, bullate leaves.

Moraceae

Maquira guianensis

Naucleopsis naga

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Moraceae

Perebea xanthochyma

Perebea xanthochyma

Perebea xanthochyma (cerillo). A medium-sized tree of forest understory. The trunk is typically irregular and has long, hanging branches. Leaves are fairly large, simple, alternate, toothed, with pubescence; branchlets are also pubescent. Tertiary veins are ladderlike: densely packed and parallel. The stipule encircles the branchlet at the tip and leaves behind a clear ring scar after it falls. Any broken part produces lots of white latex. Flowers are tiny, white, in single little balls held at the base of each leaf. Fruits be-

come red. d istribution: In lowland wet and sometimes lower montane forest. One of the dominant subcanopy trees of mature forest in the wetter parts of the Canal Area, especially at Pipeline Rd. Not seen in open areas or along forest edge. Recognition: Readily recognized as Moraceae by the encircling stipule scars and latex. Castilla elastica (not illustrated) is closest in overall leaf size, shape, and pubescence, but has much finer teeth and a broad leaf base. The other toothed Moraceae (Sorocea affinis and Maclura tinctoria) lack pubescence. Trophis caucana has similar-sized leaves with teeth, but they are sandpapery. Two additional species of Perebea are found in Panama, both wet-forest species. One is very rare. The other occurs in wet forest near the Canal and sparsely in Costa Rica, has teeth but no hairs, and resembles Sorocea affinis.

Poulsenia armata

Poulsenia armata

Poulsenia armata (cucua). A tall canopy tree of

moist to wet forest. Large individuals have dark, almost black bark, and large buttresses. The trunk is straight and branched only near the top. Leaves are large, rounded, and have small sharp spines along the underside of the primary vein. In saplings, leaves can be very long and broadly oval, whereas canopy leaves are much smaller and round. The end bud of each branch is covered in small, very sharp spines, and saplings have inconspicuous but very sharp

spines on the trunk. At the base of each leaf, there is a conspicuous scar that encircles the branch. Any broken part produces white latex, though not a lot. Yellow flowers are held in tiny dense clusters at the leaf bases, and they develop into small fruit clusters. d istribution: Known in moist, wet, and lower montane zones. Very common in mature forests at Barro Colorado I. and Soberania, where both saplings and canopy adults are frequent; it is particularly common on moist slopes, for instance on the slopes just above streams at Pipeline Rd. Not seen along roads or in clearings. The 1983 El Niño drought in Panama caused a large die-off of this species at Barro Colorado I., but it remains fairly numerous. Recognition: The spiny end bud in saplings is unmistakable. Large trees can usually be identified by leaves on the ground. No other species with simple, roundish leaves has tiny spines along the leaf vein.

Moraceae

Perebea xanthochyma

Poulsenia armata

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Moraceae

Sorocea affinis

Sorocea affinis

Sorocea affinis (cauchillo, lechosa, gallote, morillo, ramoncito). A widespread understory treelet. The small trunk has white lenticels. Leaves are narrow, dark green, with yellowish green veins below, and sometimes with sparse teeth. The narrow tip is very elongated and often curves to one side. Any broken part produces lots of white latex. Flowers are small and white, held loosely on a long spike. Fruits are berries carried on this spike. d istribution: All forests in Panama except the driest zone; sparser in Cos-

ta Rica. Common in most of the Canal Area and abundant at Barro Colorado I. and Soberania. Recognition: Leaves have a typical Moraceae appearance, although narrower than other species; see Brosimum alicastrum, Maquira guianensis, and Trophis racemosa, all fairly similar but easily separable upon study. Leaves of S. affinis have somewhat denser, more reticulate venation than the others. A species we do not illustrate, Pseudolmedia spuria, is very similar to Sorocea and is found widely in moist and wet forests, very commonly near the Canal. Three more Sorocea species are known in the region. S. pubivena is widely seen in wet and lower montane zones, less common than S. affinis in the Canal Area but more common in Costa Rica. Another species is from mountains in w. Panama and Costa Rica, the other sparsely distributed in e. Panama. All have narrowish leaves and extended tips, and two are welltoothed while S. pubivena lacks teeth.

Trophis caucana

Trophis caucana

Trophis caucana (lija). A small tree of roadsides and streamsides. The trunk often leans and forks near the ground. The bark is cream-colored with black lenticels. Small branchlets are pubescent and rough to the touch. Leaves are simple, alternate, large, toothed, with long pointed tips. They are ar-

ranged regularly along the branches, in a flat plane, and each leaf curves forward toward the end of the branch. The underside feels rough, like sandpaper. Stipules encircle the tip of the branch, but fall readily, leaving a circular scar around the branch at each leaf. Broken leaves produce latex, though sparse compared to other Moraceae. Flowers are in dense spherical heads, orange or greenish, produced throughout the year. Fruits are small and green with this same spherical form, and can also be found throughout the year. d istribution: In moist, wet, and lower montane forest. Very common along Pipeline Rd., and frequently occurs along forest streams. Recognition: An easy species to recognize from the large leaves that curve forward and their sandpapery underside.

Moraceae

Sorocea affinis

Trophis caucana

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Muntingiaceae Trophis racemosa

Trophis racemosa

Trophis racemosa (morillo, gallote, lechosa). A

medium-sized, very widespread tree. Leaves are simple, alternate, toothed, and have an extended, pointed tip and finely reticulate tertiary venation. There are usually a few teeth toward the end of the leaf. The leaf underside is rough to the touch. On young plants, leaves can be deeply lobed, in the form of a mitten. Broken leaves produce latex rather slowly.

d istribution: Very widespread, including dry, wet, and lower montane sites. It is in old forest at Barro Colorado I. and Soberania, but also occurs in open areas. Recognition: The leaves are similar to those of Sorocea, and see also Maquira. Juvenile leaves might be confused with those of Dendropanax arboreus (Araliaceae), which are also irregularly lobed but thicker and without latex. Panama has two more species of Trophis, found in mountains of w. Panama through Costa Rica. We excluded five genera in the Moraceae. We briefly mentioned Pseudolmedia spuria, and there are two other Pseudolmedia species, plus Clarisia biflora, and there is a second Clarisia. The other genera are Batocarpus (two species), Helicostylis (one), and Morus (one). The last is the temperate mulberry genus, and occurs only at high elevation in w. Panama and Costa Rica. Helicostylis is also found in mountains, but the others are lowland genera.

Muntingia calabura

Muntingiaceae Muntingia calabura

A tiny family of the American tropics, with just three species. It sometimes gets merged into the Malvaceae. We cover just one of two species known in Panama.

Muntingia calabura (periquito, pacito, majaguil-

lo, capulín). A small, weedy tree of wasteland. Leaves are toothed, alternate, regularly spaced along branches, in a flat plane; adjacent leaves overlap. The

leaf base is highly asymmetric, and the leaf underside is a light, bluish green. Three main veins arise together from the leaf base. The pretty, white flowers are solitary, held on branches at the petioles, and nearly always present. Fruits are small berries that turn red when mature. d istribution: Widespread, mostly on the Pacific coast. Only in grasslands, never in forest, not even secondary forest. Recognition: Leaves are quite similar to those of Guazuma (Malvaceae—Byttnerioideae), Trichospermum (Malvaceae—Grewioideae), and Trema (Cannabaceae). But Muntingia has softer, highly asymmetric leaves, and nearly always has white flowers. One other genus, Dicraspidia, is known in Panama. It is found throughout Costa Rica but is known only in far e. Panama, and is very similar to Muntingia but with larger, yellow flowers.

Muntingiaceae

Trophis racemosa

Muntingia calabura

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Myricaceae

Myricaceae Morella cerifera

Morella cerifera

50 species, with 12 in tropical America. Panama has just one species of montane forest, and we thus omit a family description.

Morella cerifera. A treelet of cloud forest. Leaves

A small family of treelets and shrubs mostly of temperate and subtropical areas, but with a number of species occurring in tropical mountains. Several species are known as wax myrtles (genus Myrica) for wax collected off the berries, and are found in both Europe and North America. Globally there are about

are simple, alternate, narrow and pointed, densely clustered at the ends of branches, and have large, widely spaced teeth. Male flowers are yellowish to red, on long spikes held at the petiole base. Female flowers are on spherical heads. Fruits are round berries with a whitish, waxy coat. d istribution: In lower mountains, from c. Panama through Costa Rica. Fairly common in open areas at Cerro Campana. Recognition: The teeth and bunched leaves are very distinctive. Hedyosmum (Chloranthaceae) is another montane genus with teeth, but has opposite leaves.

Compsoneura capitellata

Myristicaceae Compsoneura capitellata

This is an important tropical tree family, with most forests having several to many common and conspicuous species. It is only modest in terms of species numbers, with 400 species worldwide and 84 in the American tropics, but nearly all are large trees, and they are all tropical. This is the family of nutmeg (Myristica fragrans), a species from Southeast Asia. Panama has 19 species, and we cover 8 and describe 1 more. Myristicaceae all look similar to one another, so it is an easy family to characterize and to learn. The most obvious trait is the branching pattern, which even takes the name myristicaceous. Branches emerge in groups of three or four from the main stem or trunk, exactly horizontal and perpendicular to the trunk (verticillate); successive sets of branches are spaced at roughly equal intervals up the trunk. The simple, untoothed leaves are regularly spaced and arranged in a flat plane along the branches. Many Myristicaceae leaves have pubescence on the leaf underside, imparting a lighter tint, sometimes bluish

or whitish, but many species lack this trait. Also, Myristicaceae produce clear but reddish latex (the origin of many local Spanish names with “sangre”). Overall, the leaf pattern resembles that of Annonaceae, but Annonaceae lack latex and have branches that zigzag between leaves. Flowers of Myristicaceae are small and nondescript, but fruits of Myristicaceae are distinct, consisting of a single big seed with a bright-colored and edible sheet called an aril around it. In many species, the aril is composed of wrapping fingers that leave part of the seed visible. These seeds are held in dry capsules that split open on the tree, allowing birds or monkeys to get at the aril.

Compsoneura capitellata (sangrilillo, miguela-

rio, velario). A medium-sized wet-forest tree. Branches are verticillate. Leaves are simple, alternate, regularly spaced, but lack any pubescence below. Veins on the underside are elevated, and a very clear vein along the margin connects secondary veins; tertiary veins are closely parallel and finely spaced. Any broken part produces a flow of clear latex that quickly turns reddish. Seeds are completely enclosed in a red aril. d istribution: Only in wet forests of c. Panama, but common there. Widespread at Santa Rita, Cerro Jefe, and the upper Chagres basin. Recognition: The genus Compsoneura lacks the pubescence that helps characterize Virola and Otoba, and should be reminiscent of Annonaceae. Leaves also resemble those of Maquira guianensis (Moraceae). The red sap and strong marginal vein are cues for this species.

Myristicaceae

Morella cerifera

Compsoneura capitellata

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Myristicaceae

Compsoneura sprucei

Compsoneura sprucei

Compsoneura sprucei (sangrilillo, miguelario, velario). An understory treelet of the wet zone. Branch-

es are verticillate and sap reddish. Leaves are simple, alternate, regularly spaced, with neither pubescence nor a marginal collecting vein. Tertiary veins are closely parallel and finely spaced. d istribution: In wet forests along the Caribbean slope in Panama; widespread in Costa Rica. Recognition: Venation is much different than in C. capitellata, but both species lack pubescence on the leaf underside. This species is reminiscent of Guatteria (Annonaceae). A third, very rare, Compsoneura species is known from only a few records in e. Panama.

Otoba acuminata

Otoba acuminata

Otoba acuminata (sangrillo de montaña, miguelario, velario, saba). A medium-sized tree of mature wet forests. Leaves are simple, alternate, pointed at the tip, and conspicuously whitish gray on the underside due to a dense coat of hairs. Broken parts ex-

ude a reddish sap. The seed has a white aril. d istribution: Quite common in wet and montane forests of the Canal Area; otherwise known rather sparsely at wet and lower montane sites. Recognition: Leaf undersides are obvious even in canopy trees. O. novogranatensis is similar but has wide, rounded leaves with darkish lines below. Most of the Virola species have leaf undersides with a bluish hue, but V. elongata (not illustrated) is whitish and might be confused with Otoba. The other species with so much white on the leaf underside is Licania hypoleuca (Chrysobalanaceae), but its leaves are smaller. Otoba seeds have white arils, while other genera in the Myristicaceae have red arils.

Myristicaceae

Compsoneura sprucei

Otoba acuminata

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Myristicaceae

Otoba novogranatensis

Otoba novogranatensis

Otoba novogranatensis (miguelario, velario,

sangrillo de montaña, fruta dorada). A large wet-for-

est tree. Similar to O. acuminata, but this is a tall tree with large buttresses. The leaves also differ: in O. no-

vogranatensis they are usually blunt-tipped and the undersides have vague dark lines running parallel to the midvein; these can usually be seen even in canopy trees. d istribution: Caribbean wet forest and montane areas throughout. Common in Bocas del Toro. Recognition: See O. acuminata. Leaves also resemble those of Parathesis amplifolia (Myrsinaceae), which is a smaller tree with browish, not whitish, leaf undersides. Two more Otoba species are found in the region; both are wet-forest specialists but seen much less often.

Virola koschnyi

Virola koschnyi

Virola koschnyi (fruta dorada, sangrillo). A tall

wet-forest tree. Its buttresses can be large, and branches are high, emerging perpendicular to the

trunk in groups of four. Twigs have red pubescence. Leaves are very large, alternate, regularly spaced, in a flat plane, light greenish to bluish below with contrasting red veins covered in pubescence. There are many secondary veins, parallel and closely spaced. Broken leaves exude a clear sap that turns reddish. d istribution: Wet lowland forests throughout. Recognition: Virola are fairly easy to recognize from the light leaf undersides with a bluish tint and the usual leaf and branching patterns of Myristicaceae. This species has much larger leaves than other Virola.

Myristicaceae

Otoba novogranatensis

Virola koschnyi

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Myristicaceae Virola macrocarpa

Virola macrocarpa

Virola macrocarpa (velario, miguelario). A tall

Branching is verticillate, and leaves are alternate, in a flat plane. Leaves are very light below, with a greenish or bluish tint; veins are yellow. Produces reddish sap from broken parts. d istribution: Primarily known in wet and montane forests east of the Canal, but not very common; we also found it in Bocas del Toro, and it is known in s. Costa Rica. Recognition: The leaf is shaped like that of V. sebifera, but lacks the reddish pubescence. See also O. acuminata, which has a whitish or cream color to leaf undersides.

wet-forest tree. On big trees, bark splits into plates. Virola multiflora

Virola multiflora

Virola multiflora. Not illustrated. A tall forest tree. Like other Virola in all respects, but the leaves are very narrow and pointed. They have the characteristic bluish tint below. d istribution: A Caribbean wet-forest species, but it makes it to Barro Colorado I. and Soberania, where it is fairly common. Recognition: Big trees are easy to spot when the verticillate branching is visible, because the narrow, pointed leaves are distinctive even from far away.

Virola sebifera

ish with contrasting red veins, all due to pubescence. Broken leaves produce sap that turns reddish. d istribution: In forests everywhere except the driest stretch of the Pacific coast, often very common. Found everywhere in the Canal Area, including forest edge, secondary forest, and old growth, but does not grow in open areas. As in all the Virola, saplings are common in the shade. Recognition: This is the abundant Virola, easy to learn because it is seen so often. In Pacific regions, for example around Panama City, it is the only Virola, but it also occurs in wet forests with V. macrocarpa and other similar species. Check also the leaf undersides of Mosannona garwoodii (Annonaceae), which are much bluer than in Virola.

Virola sebifera

Virola sebifera (velario colorado, copidijo, boga-

mani, sangre). A very widespread medium-sized tree. Branching is conspicuously verticillate. Twigs have red pubescence. Leaves are alternate, regularly spaced, and in a flat plane. The leaf underside is blu-

Myristicaceae

Virola macrocarpa

Virola sebifera

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Myrsinaceae

Virola surinamensis

Virola surinamensis

Virola surinamensis (fruta dorada, miguelario, bogamani). A tall wet-forest tree. Buttresses can extend far up the trunk. Branches are horizontal, in groups of three or four. Leaves are alternate, regularly spaced, in a flat plane, long and straight-sided. The widest point of the leaf is near the tip. Undersides are light greenish blue, and the veins are yellow. Broken leaves produce reddish exudate. d istribution: In Caribbean wet forests throughout Panama, reaching Barro Colorado I. and Soberania in moister valleys,

but not very common. In Costa Rica known only at a few sites. Recognition: The long leaves that are widest beyond the middle set it apart from other Virola except V. koschnyi, but leaves of V. koschnyi are considerably larger and wider, with reddish veins. Three more Virola species are known in Panama. We mentioned V. elongata under Otoba: it is common in Canal Area wet forests but nowhere else, and looks like V. sebifera but with whitish leaf undersides. V. guatemalensis is rare in Panama but common in mountains and wet forests of Costa Rica. The last is seldom seen. We have not yet mentioned two other genera of Panama’s Myristicaceae. Iryanthera has three species in wet forests and rather resembles Compsoneura but has pubescence like that of Virola and Otoba. The other genus, Osteophloeum, has just one species, and its leaves narrow to a pointed base, unlike the rest of the family.

Myrsinaceae Ardisia bartlettii

Ardisia bartlettii

A large tropical family, almost entirely shrubs but some treelets or small trees, with 1250 species worldwide and 400 in the American tropics. This is the sixth largest family among Panama’s trees and treelets with 75 species, tied with the Annonaceae. Because Myrsinaceae are all small and uncommon in lowland forests, they are not considered an important component of rainforest vegetation. At high elevations, though, they can be prominent. We cover six of Panama’s species here. All Myrsinaceae have simple, alternate leaves, rather nondescript except for the presence of tiny, translucent dots or lines in the leaves, visible with magnification while holding a leaf against bright light. These “punctuations” are subtle, and it takes some practice, but it is a good character. Both Salicaceae and Myrtaceae are known for similar leaf dots, but the former usually have teeth and the latter opposite leaves, and Myrsinaceae is the only family that sometimes has dark, translucent lines. In many Myrsinaceae, leaves are thick, stiff, and shiny, with inconspicuous venation, though many other tree families have leaves like this. Indeed, these thick leaves look like they ought to have latex, but do not

(leaves with latex are usually thickened and stiff). Flowers and fruits are small, in terminal clusters, and not especially distinctive.

Genus Ardisia. This is the third largest genus in Panama’s checklist of trees and treelets, with 54 species (Psychotria also has 54, Inga has 55, and Miconia has 65). Moreover, there are 39 other Ardisia that are too small to be counted as treelets, and this division is imprecise, so you might want to keep in mind that there are 93 species of the genus in Panama. An entire book would be needed to cover them. Most Ardisia are montane, and many are in wet forests. Here we illustrate three species and mention a fourth that occur in lowland forests. In montane or Caribbean wet forests, you may have to let many Myrsinaceae go as unknown Ardisia. Ardisia bartlettii (canelito, canelo). A treelet of

forest understory. The small stem typically has low branches. Leaves are simple, alternate, shiny and rather thick, sometimes slightly toothed though often not at all. Held up to bright light and studied with a hand lens, leaves show clear spots mixed with short black lines. Flowers are small and pink, and their sepals (the floral appendage below the petal) have the same pellucid spots and lines that leaves have. Fruits are small berries. d istribution: Known mainly from the Caribbean slope of the Canal east, including Barro Colorado I., where it is very rare. Recognition: Thick and shiny leaves can be learned if you remember to check for the pellucid dots, which are distinctive. Of the Ardisia we cover, this is the one with teeth.

Myrsinaceae

Virola surinamensis

Ardisia bartlettii

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Myrsinaceae

Ardisia revoluta

Ardisia revoluta

Ardisia revoluta (canelito, canelo). A dry-forest treelet. Leaves are smooth, shiny, rounded at the end, and secondary veins are barely visible. Flowers are white. d istribution: Of common Ardisia, this is the only one in dry areas. It is quite common in riverine forests of the Pacific slope of c. Panama and nw. Costa Rica. Recognition: Other Ardisia do not grow in the dry zone. Stylogyne also occurs in forests near Panama City, but has narrower, pointed leaves.

Ardisia standleyana

d istribution: The commonest Ardisia at Barro Colorado I., though still fairly rare. Also seen in Soberania and the Caribbean coast near the Canal and sparsley through Costa Rica. Recognition: Leaves are quite similar to those of Aspidosperma spruceanum (Apocynaceae), but Aspidosperma has latex.

Ardisia standleyana

Ardisia standleyana (canelito, canelo). A treelet

of forest understory. Much like the other Ardisia, with no teeth on the leaves and white or pink flowers.

One more Ardisia species is found on Barro Colorado I., A. guianensis (not illustrated). It has almost invisible secondary veins, like those of A. revoluta. Many of the other Ardisia are seldom seen and poorly known.

Myrsinaceae

Ardisia revoluta

Ardisia standleyana

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Myrsinaceae

Myrsine coriacea

Myrsine coriacea

Myrsine coriacea (mangle de montaña, mangle, mangle de sabanas). A small cloud-forest tree. The trunk is straight and unbranched for much of its height. Bark is white or gray with lenticels. Twigs have reddish hairs. The simple, alternate leaves are

narrow, pointed, bunched toward the ends of branches, and have tiny translucent dots and purplish lines. Flowers are green or white and produced along the branches. d istribution: Common at Cerro Campana and known in mountains of w. Panama. Widespread through mountains of Costa Rica. Recognition: Unlike Ardisia, this is a real tree, though not a large one. Resembles Morella cerifera (Myricaceae) of the same habitat, but Morella has toothed leaves. The tiny purple rays offer an excellent character, but they will not be seen without magnification. Panama has one more species of Myrsine. It has larger leaves, but is rarely seen.

Parathesis amplifolia

Parathesis amplifolia

Parathesis amplifolia. A small tree of wet and montane forests. Branchlets have fine reddish pubescence. Leaves are simple, alternate, brown on the underside from a dense coat of hairs, with translucent dots. There is a slightly darkened band running parallel to the central vein. Flowers are white.

d istribution: Wet Caribbean and lower montane forests. Easy to find at Cerro Jefe and Cerro Campana and environs. Not known in Costa Rica. Recognition: The brown leaf undersides distinguish it from other Myrsinaceae. See species of Otoba (Myristicaceae), which have leaves that are whitish or brownish below, especially O. novogranatensis, which has a dark band along the midvein. Chrysophyllum cainito (Sapotaceae) and Luehea seemannii (Malvaceae—Grewioideae) also have brown leaf undersides, but they should not be confused. Nine more Parathesis species are known in Panama, most being small trees like P. amplifolia. Not all have brown leaf undersides. Two are restricted to montane forest; others are seldom seen.

Myrsinaceae

Myrsine coriacea

Parathesis amplifolia

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Myrtaceae

Stylogyne turbacensis

Stylogyne turbacensis

Stylogyne turbacensis (canelito). A small forest tree. Leaves are simple, alternate, shiny and rather thick, narrow and pointed, bunched toward the ends of branches, and have translucent dots visible with magnification. d istribution: Common at Barro Colorado I. (the only common Myrsinaceae there) and throughout forests on the Pacific half of the Canal Area, plus on limestone sites near the Caribbean.

Widely but sparsley known in Costa Rica. Recognition: The easiest-to-learn lowland Myrsinaceae, and the only one routinely encountered near the Canal. The narrow, bunched leaves are quite different from other Myrsinaceae, rather resembling those of Thevetia (Apocynaceae) and Tovomita species (Clusiaceae), but both of those have latex. Two more Stylogyne species are found in Panama, but they are seldom seen. In addition, there are four genera of Myrsinaceae we have not mentioned: Cybianthus and Hymenandra with two species each, and Geissanthus and Gentlea with one. Cybianthus schlimii is common at Cerro Campana and through mountains of Costa Rica. Hymenandra is tree-sized and known in lower montane and Caribbean wet forests of Panama and Costa Rica. The other two genera are seldom collected.

Myrtaceae

Calycolpus warszewiczianus

Calycolpus warszewiczianus

The myrtle family is one of the dominant families throughout the world’s tropics and subtropics, plus warm temperate Australia. There are about 3500 species worldwide and 1500 in tropical America; nearly all are trees, but there are a few shrubs. This is the family of Eucalyptus, making it the dominant family across most of Australia, and it also includes wellknown species such as the clove tree (Syzygium aromaticum) of Asia, allspice (Pimenta dioica) of Central America and Mexico, and guava (Psidium guajava). In Panama, there are 45 tree species in the Myrtaceae, and we cover 8 native species plus 1 exotic. As a family, the Myrtaceae are pretty easy to learn, but species within the family are notoriously difficult to identify. All Myrtaceae have opposite leaves, and they are not difficult to separate from the other two important opposite-leaved families: Melastomataceae, which have very distinct venation, and Rubiaceae, which have an interpetiolar stipule. Two other opposite-leaved families, Clusiaceae and Apocynaceae, have latex. Thus, Myrtaceae are most often confused with several minor families with opposite leaves, especially Vochysiaceae and Lythraceae. Generally, if you can eliminate Melastomataceae and Rubiaceae, then Myrtaceae is a good guess for any tree with opposite leaves in the American trop-

ics. There is a good character too: most Myrtaceae have a vein running along the leaf margin called a collecting vein. Be cautious, however, because other families have this trait. Another widespread trait in Myrtaceae is bark that peels off to leave a naked trunk, but only some species have this. Also, in some Myrtaceae, crushed leaves have a strong spicy smell (allspice and Eucalyptus are conspicuous examples), but in many species the odor is weak or absent. Finally, Myrtaceae leaves should have clear, tiny dots, visible only when held up to the light and with magnification; these translucid spots, though, are not easy to see. Flowers are typically recognizable; they are flat, with white petals and many stamens, and usually fairly large. Fruits are fleshy spheres, and many are soft, sweet, and edible.

Calycolpus warszewiczianus (guayabillo, guay-

abito de monte). A small tree of dry areas. The bark is brown or gray, and peels away to reveal white and smooth inner bark. Leaves are simple, opposite, smooth and shiny green, with closely spaced, parallel secondary veins and a distinct collecting vein. Translucid points are visible with magnification. Flowers are large, white to pink, with a dense mass of pinkish stamens. d istribution: Known mostly in c. Panama, and there on the Pacific slope, including agricultural areas. We have also found it on the Caribbean slope of the Canal Area, and there are records in s. Costa Rica. Recognition: The smooth, shiny leaves resemble those of Myrcia gatunensis. The trunk is like that of the guava (Psidium guajava) or Eucalyptus, but neither of those has smooth, shiny leaves. Few Myrtaceae have pink flowers, and Calycolpus can have flowers at any time of year.

Myrtaceae

Stylogyne turbacensis

Calycolpus warszewiczianus

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Myrtaceae

Eugenia coloradoensis

Eugenia coloradoensis

Eugenia coloradoensis (guayabito, guayabito

de montaña). A medium-sized forest tree. The trunk

is straight, cylindrical, and the bark brown or reddish. Leaves are opposite, in a flat plane, smooth, with a weak collecting vein. Translucid points are visible with magnification. The small fruits mature black or purple. Distribution: Widespread near the Canal, in all types of forest; otherwise seldom seen. Recognition: Easily recognized as Myrtaceae. This species has somewhat larger and wider leaves, lacks wavy margins, compared to other Eugenia and Myrcia. Check E. nesiotica. Another Eugenia of the Canal Area, E. galalonensis (not illustrated), is also similar.

Eugenia nesiotica

Eugenia nesiotica

Eugenia nesiotica (sequara). A medium-sized forest tree. The trunk is ribbed. Leaves are small, opposite, in a flat plane, with wavy margins, smooth, and

shiny green. Collecting veins arch from one intersecting vein to the next. Translucid points are visible with magnification. Distribution: Almost entirely restricted to forests around Barro Colorado I., but it is rather common there. Recognition: The wavy leaf margins are close to those of E. oerstediana. It is also very similar to Mouriri myrtilloides (Melastomataceae), and student botanists at Barro Colorado I. are usually quizzed at some point with the near-identical leaves of the two species side-by-side. In Mouriri, there is no petiole. Lafoensia punicifolia (Lythraceae) is also similar, but its leaves have a small pit near the tip.

Eugenia oerstediana

Eugenia oerstediana

Eugenia oerstediana (guayabillo). Not illustrated. A medium-sized forest tree. The leaves are small, opposite, similar to those of E. nesiotica but with more pronounced venation. Translucid points are visible with magnification. Distribution: Widespread in moist to wet lowlands and some lower montane sites. Common in old growth at Barro Colorado I. Recognition: See E. nesiotica.

Myrtaceae

Eugenia coloradoensis

Eugenia nesiotica

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Myrtaceae

Eugenia principium

Eugenia principium

Eugenia principium (guayabillo). A small tree of

dry areas. Bark on bigger trees flakes off, revealing reddish inner bark. Leaves are simple, opposite, small and pointed, shiny green above and light green or even yellowish below. The veins are very inconspicuous and the petiole short. Distribution: Mostly

in dry areas along the Pacific coast, and abundant in forests near Panama City and on limestone outcrops near the Canal. Surprisingly, it also shows up in very wet forests in c. Panama. Recognition: Leaves are like those of Myrcia gatunensis in having inconspicuous venation, but are the size and shape of leaves of other Eugenia. Fourteen more Eugenia species are found in Panama, enough to make the genus dangerous to identify; experienced botanists have a difficult time with it. We have covered the common species near the Canal. Worldwide, Eugenia is a very large genus, and in South America, if you find a Myrtaceae you cannot identify, Eugenia is always the best guess.

Myrcia gatunensis

Myrcia gatunensis

Myrcia gatunensis (pimiento). A medium-sized

forest tree. The trunk is straight, light gray or even whitish, and branched at a low height. Leaves are simple, opposite, narrow and pointed, shiny green and smooth. Veins are inconspicuous, but close inspection shows a neat and straight collecting vein. Secondary veins are dense, straight, and run tightly parallel. Translucid points are visible with magnification. Crushed leaves can have a fairly strong, Eucalyptus-like odor. Flowers are white, and have many stamens extending well outside the petals, so they look

mostly like balls of stamens. Distribution: Widespread in all types of forest, from dry to wet and lower montane. Undoubtedly an indication of how difficult Myrtaceae are, we found only two specimens of this species in published herbarium records, yet we see it in nearly every forest we visit in Panama. Recognition: Easy to spot as Myrtaceae with opposite leaves and the collecting vein. The parallel secondary venation is much different from that of Eugenia. Twelve more Myrcia species are found in Panama, and all but M. splendens of the mountains are rarely seen. Another one in the Canal Area, M. fosteri, was named for our mentor Robin Foster, and closely resembles M. gatunensis except for the flowers. A common one at Pipeline Rd. is M. zetekiana, with leaves longer and narrower than those of any other Myrtaceae we have described. In general, there is no good way to distinguish Myricia from Eugenia without studying flowers, and even botanists have trouble.

Myrtaceae

Eugenia principium

Myrcia gatunensis

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Myrtaceae

Psidium guajava

Psidium guajava

Psidium guajava (guava, guayaba). A small, familiar tree of farmland. The small, forked or leaning

trunk loses its outer bark, leaving behind smooth, light-gray inner bark. Leaves have prominent, elevated veins below, with pubescence. Flowers are white with many long stamens. Fruits are spherical, green initially but turning yellow. Distribution: Only seen around human habitation, in yards and gardens, never in the forest. Its natural range is in the Americas but has long been obscured by cultivation. Recognition: Best known from the smooth trunk, opposite leaves, and habitat. Everyone in the tropics knows the fruit. See P. guineense.

Psidium guineense

Psidium guineense

Psidium guineense (guayabita sabanera, guayabo de sabana). A savanna treelet. The outer bark is brown but peels off in sheets. Branchlets are hairy. Leaves are simple, opposite, rounded at both ends, densely pubescent with whitish hairs, with translucent spots. Distribution: Rural areas of the Pacific slope and up into lower mountains. Not in forest. Recognition: Resembles the cultivated guava, P. guajava, the latter having narrower leaves. Densely hairy, opposite leaves might be confused with the nance (Byrsonima crassifolia [Malpighiaceae]), which grows in the same habitat but does not have smooth, gray, peeling bark. Two more Psidium species are found in Panama. P. friedrichsthalianum is a forest species, known as a very rare tree at Barro Colorado I., Gamboa, and a few sites in Costa Rica. It has leaves much like those of Eugenia or Myrcia but smooth, peeling bark like that of the guava. The other species is known from only a few sites in the mountains.

Syzygium malaccense (marañón curasao, roseapple). Not illustrated. A large tree imported from Asia. Leaves are opposite, shiny, densely concentrated on branches, forming a thick crown. The flowers are large and pom-pom–like, a deep rose color, produced on large branches or directly from the trunk. Fruits are red on the outside but white within, soft and edible but often with a hint of bitterness. Distribution: Widely planted in towns and yards. Never in forest. Recognition: Best known from its flowers and fruits. The Panama checklist has a second widely planted Syzygium, S. jambos, also from Asia. Myrtaceae has six native genera we have not yet mentioned: Calyptranthes, Chamguava, Myrcianthes, Myrciaria, Plinia, and Siphoneugena. Calyptranthes is a big genus, with nine species, but all are from wet or montane forest and most are rare. Myrciaria and Myrcianthes have two species each, and the other genera just one. The Chamguava species is found at Barro Colorado I., though not common. Myrtaceae species and even genera are difficult to separate from one another; many have fairly similar leaves. Particularly away from the Canal Area, and in wet or montane forests, you will have to let many go as unidentified. To complete a discussion of Myrtaceae, we also have to mention Eucalyptus, which is widely planted in Panama and all warm parts of the world. Panama’s checklist includes six species, all imported from Australia, but we will not take space here to consider how they are identified to species.

Myrtaceae

Psidium guajava

Psidium guineense

357

358

Nyctaginaceae

Nyctaginaceae Guapira standleyana

Guapira standleyana

A moderate-sized family that mixes herbs, shrubs, trees, and lianas of both tropical forest and subtropical deserts. There are 400 species worldwide and 180 in tropical America. Included are two familiar garden plants, the four-o’clock (Mirabilis, from North American deserts) and Bougainvillea (from dry parts of South America). There are eight species in Panama’s tree checklist, and we cover two. The tropical forest Nyctaginaceae all have simple, opposite leaves that are crowded at branch tips and sometimes even whorled (several leaves emerging from a single point on a branch). Because they lack stipules, they can be distinguished from Rubiaceae, which as a family is far more common. Malpighiaceae also have opposite, crowded leaves, but they too have stipules. Flowers of the Nyctaginaceae lack petals, but are small and not particularly distinctive. This is an inconspicuous family, mostly shrubs or treelets that are seldom common, and thus not easily learned.

Guapira standleyana (mala sombra, malvecino, llanto). Having just commented that most are small

treelets, we start with the one large forest tree. The trunk is light brown, strongly ribbed, and spreading into small buttresses at the base. Leaves are opposite, strongly crowded toward the tips of branches, smooth and bright green; there is pubescence on both leaves and twigs. Flowers are tiny, yellow to green, with stamens sticking out but no petals, in little clusters of clusters on long stalks emerging from the leaf nodes. Fruits are fleshy spheres that turn purple when mature. Distribution: We record it regularly in mature forest at Barro Colorado I. and Soberania, but it is seldom seen elsewhere and not known in Costa Rica. Recognition: When large, it is readily recognized from the fluted trunk. It can be confused with Quararibea asterolepis (Malvaceae—Bombacoideae), but Quararibea has peeling bark and Guapira does not. As a sapling, Guapira does not stand out. Look closely at the bunched leaves to see that they are opposite but lack stipules, setting it apart from most similar-looking leaves. Byrsonima species (Malpighiaceae) have leaves in a similar arrangement, but quite different in appearance, and with stipules. See also Terminalia (Combretaceae) with bunched leaves, though they are alternate; however, in both Guapira and Terminalia, leaves can be so bunched that it is difficult to tell whether they are alternate or opposite. One other Guapira species is widespread but rare. It has similar bunched, opposite leaves, but the trunk is not fluted.

Neea amplifolia

Neea amplifolia

Neea amplifolia (canelo, canelito). A small, un-

derstory treelet. The slender stem branches at a low height and often leans. Leaves are simple, opposite, rather thick and shiny green, and bunched toward the tips of branches. Twigs have dense red hairs near the tip (among the leaves), and leaves also have some red pubescence on veins on the underside. Flowers are pink, in little clusters, and lack petals.

Fruits are berries. Distribution: Found widely in lowland moist and wet forest, mostly on the Caribbean slope, probably overlooked. Recognition: An inconspicuous little treelet, Neea can be confused with Psychotria (Rubiaceae), most of which are also understory shrubs with opposite leaves; another Rubiaceae, Hamelia patens, has leaves whorled at the branch end. But all Rubiaceae have stipules, which all Neea lack. Five more Neea species are found in Panama. All are similar to N. amplifolia: small plants that barely pass as treelets. N. delicatula is fairly common near the Canal, even in areas around Panama City, but rare elsewhere. There are a couple montane species common in Costa Rica but rare in Panama, and two others very poorly known.

Nyctaginaceae

Guapira standleyana

Neea amplifolia

359

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Ochnaceae

Ochnaceae

Cespedesia spathulata

Cespedesia spathulata

A moderate-sized tropical family, mostly shrubs but also herbs and trees, with 500 species worldwide and 300 in the Americas. There are 13 species in Panama, of which we cover 2. The family does not characterize well. Most have simple, alternate, toothed leaves and dark brown stipules. In other ways, the genera in Panama are so different that they must be learned separately.

Cespedesia spathulata (membrillo de montaña,

membrillo macho, candelabro). A medium-sized tree with very large leaves. The trunk can be unbranched, or have a few large branches that rise vertically (like additional trunks). Leaves are close to 1 m long, narrow at the base, toothed, with a broad, rounded tip. They are tightly clustered at the top of the trunk, in palmlike style. The leaf cluster is held

precisely, with the upper leaves pointing upward, the lower ones downward; the leaves do not droop. New leaves are so red that they look like flowers at a distance, but they quickly turn green. Look closely at the leaf underside: the tertiary veins are neatly arranged and closely parallel. Flowers are brilliant yellow, and are held in spikes on top of the leaves. Distribution: In wet and lower montane forests throughout. In c. Panama, it is very common along roadsides, and its abundance can be used as a guide to the moister climates. It is common along the outer half of Pipeline Rd. and other forested roads near the Caribbean, scarce around Gamboa, absent near the Pacific coast except on low mountains. It is primarily a species of forest clearings and roadsides. Inside the forest, it is rare, and it is seldom seen as a juvenile. Recognition: The large leaves clustered at the top are easy to spot, but they are very similar to those of Gustavia superba (Lecythidaceae). In fact, a paper published in the eminent journal Science in the early 1970s confused the two species. The key feature is how leaves droop in Gustavia but not in Cespedesia. When leaves can be examined closely, Gustavia lacks the fine, parallel tertiary venation visible on the underside in Cespedesia. Additionally, Gustavia lacks red leaves, stipules, and has completely different flowers.

Ouratea lucens

Ouratea lucens

Ouratea lucens (Mickey Mouse). An understory

treelet of mature forest. Leaves are simple, alternate, finely toothed, with very distinct venation: secondary veins are close together and faint, and each curves far forward, some reaching almost to the leaf tip. Flowers are bright yellow, in long spikes at the ends of branches. Fruits are formed of a red receptacle (forming from the flower stem) to which tiny black

berries adhere in groups of three; these groups look like a mouse face with two ears, hence the common name. Distribution: Very widespread in all types of forest, from dry to lower montane. Common throughout the Canal Area, but only inside the forest. Recognition: No other genus has such arching secondary veins, so Ouratea is easy to learn and not forgotten. Panama has nine more species of Ouratea, and they all have arching secondary veins, though not as extremely arching as in O. lucens. O. prominens occurs in dry areas and on limestone near the Canal, and is widely known in Costa Rica. The remaining species are restricted to wet forest or very rarely seen. One other genus of Ochnaceae, Elvasia, is known from only a handful of sites in both Panama and Costa Rica.

Ochnaceae

Cespedesia spathulata

Ouratea lucens

361

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Papaveraceae

Papaveraceae Bocconia frutescens

Bocconia frutescens

A moderate-sized family of 220 species worldwide, but almost entirely north temperate herbs, including poppies. Three genera and 18 species occur in the American tropics, mostly in Central America and the Caribbean area, but several into South America as well. They are mostly found at high elevation, but not exclusively, and can be seen in wet lowland forests. Panama has just a single species of treelet (plus one

Pellicieraceae

herb species). Poppies are generally recognizable from orange latex and deeply lobed leaves as herbs, and the one treelet species in Panama shares these traits. On the other hand, poppies are best known for their brilliant showy flowers, yet Panama’s treelet species has tiny flowers without petals.

Bocconia frutescens (tabaquillo, sangrillo). A weedy, montane treelet. Leaves are toothed and deeply lobed like those of dandelions (Taraxacum officinale), with a bluish tint to the underside. They are bunched toward the top of the stem. Flowers are tiny, on a stalk above the leaves. Distribution: Grows above 1000 m elevation in c. and w. Panama through Costa Rica, and known in e. Panama. There are rare lowland records. Recognition: Watch for weedy Asteraceae with lobed leaves, but those growing as little trees are quite different.

Pelliciera rhizophorae

(or Tetrameristaceae)

Pelliciera rhizophorae

The Pellicieraceae is just a single species of mangrove, but it has been merged with two other distant species—one in Venezuela and one in Malaysia— into the Tetrameristaceae. They were all once considered part of the tea family, Theaceae. This is the only family we describe without any illustration: there is just the one species, and it easy to recognize when encountered.

Phyllanthaceae

Pelliciera rhizophorae (mangle piñuelo, pie de

santo, tea mangrove). Not illustrated. A true mangrove, found only in saltwater swamps. The base of the trunk widens greatly and has narrow, fluted ridges. The leaves are alternate, densely clustered at the ends of branches, shiny smooth green, with no venation visible. Flowers are striking: large, with five narrow, long white petals. Distribution: Only in Pacific coast mangrove swamps, not the Caribbean coast and never seen away from salt water. Recognition: Should not be confused, as Pelliciera is unlike the other mangrove species with which it associates (Rhizophora in the Rhizophoraceae, Avicennia in the Acanthaceae, and Laguncularia in the Combretaceae). Easily recognized immediately by the trunk.

Amanoa guianensis

(Formerly Euphorbiaceae)

Amanoa guianensis

are a few trees in Phyllanthus and other genera. Panama has 10 tree species, of which we cover 4. The family was once considered part of the Euphorbiaceae, but even this segregated subset of that large and variable family does not characterize well. Leaves are always simple, alternate, and untoothed, without latex. The traits for recognizing the species we cover vary and must be learned individually.

Amanoa guianensis (pantano). A medium-sized A large tropical family, with a few temperate species as well. There are 1745 species worldwide and about 300 in tropical America. Most of the family is herbaceous and in the single genus Phyllanthus, but there

tree of Caribbean swamps. The trunk has a dense mass of stilt-roots emerging near the base. Leaves are simple, alternate, short, wide, and rounded, much lighter in color below than above. At the ends of

Phyllanthaceae

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Bocconia frutescens

Amanoa guianensis

branchlets are small brown stipules. Flowers are in short clusters held along a terminal spike. Fruits are dry capsules that split open. Distribution: Known in swamps on the Caribbean coast through Panama and Costa Rica. Common near the water at the mouth of the Chagres River near the Canal. Recognition: The dense stilt-roots are bizarre, resembling only the palm Iriartea deltoidea (Arecaceae). By leaf

alone, this can be confused with a number of other species. Pera arborea (Euphorbiaceae) is quite similar, as is Bourreria costaricensis (Boraginaceae), but neither has stilt-roots or grows in swamps. The Bocas del Toro swamp specialist, Campnosperma panamense (Anacardiaceae; not illustrated), might be confused until you check the roots.

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Phyllanthaceae

Hieronyma alchorneoides

Hieronyma alchorneoides

Hieronyma alchorneoides (zapatero, pilón, palo chancho, piedro). A tall and well-known tree. The trunk is straight and cylindrical, and the bark is light brown, granular, and peels in small pieces. Large trees have small buttresses at the base, and these extend into roots that spread on the soil surface far from the trunk. Leaves are simple, alternate, and clustered toward the end of branches; their size and shape is quite variable, but most often they are moderately wide and rounded. Secondary veins are pronounced and elevated on the leaf underside. Large trees typically have some yellow leaves in the crown. The stipules of saplings are remarkable—green and leaflike, but rolled into a little structure that ants sometimes inhabit (indeed, the stipule appears to be

designed to house small creatures). In adults, the stipules are nothing like this, instead they are thin and short. The tiny flowers are on long, weak spikes held at leaf nodes. Fruits are small berries on the same long spikes. Distribution: Widespread in all lowlands except the driest zones, sometimes in swamps. Mostly a tree of forest edge or natural clearings; within the forest, there are scattered large trees but few saplings. Despite the edge preference, not a tree of towns or open areas. Recognition: Leaves are not especially distinctive, but the combination of a bunched arrangement and roundish shape help mark this species. Similar leaves are seen in the Euphorbiaceae, where this species was formerly classified. Alchornea latifolia (a euphorb) is similar, but has slightly toothed leaves. Tetrorchidum (a euphorb) has leaves of a similar shape, but with fewer secondary veins and small glands. Hura and Croton (also euphorbs) have heart-shaped leaves. Hieronyma might be confused with Terminalia amazonia (Combretaceae), whose leaves are similar in shape and also clustered toward branch ends. Juveniles of Hieronyma are readily recognized by the bizarre stipules. Adults can be learned from the light brown bark, long roots, and swollen buttresses.

Hieronyma oblonga

Hieronyma oblonga

Hieronyma oblonga (trompito). A medium-

sized tree of cloud forest. The trunk often has low

branches and lacks buttresses. Leaves are similar in shape to those of H. alchorneoides, but are light yellowish and scaly below; branchlets are rough to the touch from scales. Stipules are small and simple, never swollen into hollow structures. Flowers and fruits are on short stalks held at leaf nodes. Distribution: A cloud-forest and lower montane species throughout, also at a few very wet lowland sites. Recognition: See H. alchorneoides, which has similar leaf form but differs in most other respects. The two species do not, by-and-large, overlap in distribution.

Phyllanthaceae

Hieronyma alchorneoides

Hieronyma oblonga

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Piperaceae

Margaritaria nobilis

Margaritaria nobilis

Margaritaria nobilis (clavito). A small tree of all

types of forest. The trunk is short and branched. Small branchlets are dark brown, with conspicuous white lenticels (little pores). Leaves are simple, alternate, small, dark green above and lighter green below. They are spaced fairly regularly along branches. Flowers are small, in little groups at branch nodes. Fruits are lobed capsules that open to reveal shiny, brilliant blue seeds. Distribution: Widespread in moist, wet, and lower montane sites. Scarce inside mature forest; more often in open areas, but not common. Recognition: Without the brilliant blue seeds,

this species is characterized only by a lack of distinctive characters. It is similar to the equally nondescript Pera arborea (Euphorbiaceae), and Drypetes standleyi (Putranjivaceae) has leaves of similar size, shape, and arrangement, though quite different in detail. Margaritaria might also be mistaken for Annonaceae (Oxandra) or Lauraceae (Ocotea), and Salicaceae (Casearia and Laetia) have similar leaves, but with teeth. The other Phyllanthaceae in Panama are in four additional genera. Astrocasia and Croizatia each have one species of small trees, and both are rare. Phyllanthus is a larger genus, mostly herbs, but there are six shrubs or small trees, and one of them, P. acuminata, is widespread in Panama and Costa Rica and commonly seen in the Canal Area. It has small leaves arranged densely along branches, looking like leaflets of a big compound leaf. Finally, Richeria has two species, both large trees with nondescript though slightly toothed leaves, widely distributed in wet and lower montane forest, but not very common.

Piper reticulatum

Piperaceae Piper reticulatum

A large family of tropical shrubs and herbs, with a few lianas and epiphytes. There are 1500–2000 species worldwide, with about 1000 in the American tropics. There are 37 treelet or shrub species in Panama, all in one genus, but there are approximately 150 more herbaceous species. We cover only one species here, partly because the Piper species are mostly small shrubs, partly because they are so easy to recognize to genus, and partly because their taxonomy is in need of work, so that many species are difficult to name. We omit a family description but instead give one for the genus.

Genus Piper. This is one of the easiest-to-recog-

nize genera in the Neotropical tree flora. Stems are weak and leaning (sometimes prostrate), and have swellings on the stem at the base of every leaf and branch; the swellings are obvious and distinctive along every branch, even after leaves have fallen. Where there are still leaves, the petiole base wraps the swollen branch like a sheath. Leaves are highly variable in shape, but no other shrubs in Panama have these stem and petiole traits. Flowers of Piper are on whitish stalks that sometimes stand up, some-

times hang down; individual flowers are no more than tiny bumps, not like flowers at all. Most Piper are shrubs, some growing on the ground, some 1–2 m tall. Here we describe one species that becomes a small tree.

Piper reticulatum (hinojo). A treelet of forest understory. The small stem usually leans and forks near the ground. Where leaves emerge (or did emerge before falling), the branches are conspicuously swollen. Leaves are large, thick, heart-shaped, with many veins arising from the base and radiating like a fan. Distribution: Widespread in moist to wet and montane areas, often in the shade inside mature and secondary forest but also along forest edges. Common and conspicuous in the Canal Area. Recognition: Even without seeing the swollen stem nodes, this species is quickly recognized by the leaves. A couple other Piper species can be confused: P. marginatum also has heart-shaped leaves and occurs widely along forest edges, but its leaves are thin and papery, unlike the thick, tough leaves of P. reticulatum. P. schiedeanum also has thin, papery, heart-shaped leaves, but is much less common. At least a dozen other shrub-sized Piper species are common in lowlands near the Canal. P. arboreum and P. aequale are both frequently seen along roads and forest edge; both have small, oval leaves. In wet forests of the Caribbean slope, the genus becomes especially diverse, and throwing in herbs plus taxonomic confusion and it would take a large book to cover them well.

Piperaceae

Margaritaria nobilis

Piper reticulatum

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368

Podocarpaceae Podocarpus guatemalensis

Podocarpaceae Podocarpus guatemalensis

This is the only group of conifers native to Panama. It is a family of trees largely of the Southern Hemisphere, especially in Australia and South America, but some species’ ranges extend north of the equator in the tropics. There are 165 species worldwide and 14 in the American tropics. Panama has four species, all in the genus Podocarpus; we cover one. Leaves are not needlelike, but they are flattened, very narrow, and pointed. They lack any veins except a midrib. As in all conifers, there is no flower but rather separate male and female cones. The female cones develop into soft fruits that attract birds.

Podocarpus guatemalensis (pinotea). A tall

and cylindrical, with black bark. Leaves are densely packed on branches, deep green, smooth and shiny, narrow and sharply pointed. The central vein is elevated above the leaf underside. Distribution: Found in lower mountains of c. Panama, including Cerro Campana, but also at lowland wet-forest sites in Panama and Costa Rica. Fairly common in the mountains but rare in the lowlands. Usually only big trees are seen, seldom saplings. Recognition: At a glance, you might not immediately realize this is a conifer, especially in big canopy trees where leaves are far above. But the narrow, densely spaced leaves should be distinctive even at a distance. Virola multiflora (Myristicaceae) and Xylopia species (Annonaceae) have leaves nearly as narrow, but not held in dense clusters and nowhere near as shiny or thick. The genus Podocarpus includes three other species in Panama. Another fairly common one is P. oleifolius, which has very similar leaves but with the central vein impressed, not elevated, on the leaf underside. It is widely distributed in higher mountains of Costa Rica and w. Panama, almost never in lowlands. The other two species are rarely collected.

tree of wet and montane forests. The trunk is straight

Coccoloba manzinellensis

Polygonaceae Coccoloba manzinellensis

character. Flowers of Polygonaceae are tiny, in long stalks, and fruits come in two varieties: either berries or winged seeds.

Coccoloba manzinellensis (hueso, uvito, uve-

This is a big family worldwide, with 1100 species and 250 in tropical America, but a substantial percentage of those are herbaceous species from North America, including buckwheats and knotweeds. As a tropical tree family, it is not especially important. Panama has 22 tree species, of which we cover 4. Most Polygonaceae (and all in Panama) share one simple trait, making it a fairly easy family to recognize. The base of the leaf petiole wraps around the twig, forming a sheath that is usually large and obvious. On older branches, the sheath falls off, leaving behind a complete ring on the branch at the base of each leaf. Many Piper species have sheaths of the same variety, but Piper should be unmistakable from swollen branch nodes. With the sheath gone, the ring left behind looks like that of Ficus (and other Moraceae), but Polygonaceae have no latex. Because most individuals of the family have some younger leaves with complete sheaths, this is a reliable

ro). A medium-sized tree of the Canal Area. The trunk often forks profusely, producing many stems at the ground. Leaves are large, alternate, irregularly arranged around branches, with prominent veins raised on the underside. The stipules at the base of each leaf form a brown sheath that wraps completely around the branch. Tiny flowers are on a very long, thin spike. Distribution: Widespread in lowland forests along the Canal, from moist to wet. Also known in e. Panama. Especially common in secondary forest, and occasionally seen at the forest edge along roads, or in towns. Recognition: Leaves resemble those of Annonaceae, and check Annona hayesii, but there are no sheaths or scars in Annonaceae. See also Moraceae, which lack sheaths but do have ring scars from stipules; they always have latex, though. There are many other Coccoloba, but the large leaves and prominent veins of this species are distinctive. See also Triplaris cumingiana, with similar leaves and sheaths, but it has a single, straight stem and peeling bark. Coccoloba manzinellensis should come to mind when encountering any tree with many stems (but is not a palm) at Barro Colorado I. or Soberania.

Polygonaceae

Podocarpus guatemalensis

Coccoloba manzinellensis

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Polygonaceae

Coccoloba uvifera

Coccoloba uvifera

Coccoloba uvifera (sea grape, uvito de playa, uvero, uvito). A medium-sized beach tree. The trunk is irregular and much-branched. Leaves are alternate, wide, almost round, and the primary vein is reddish at the base. Stipules are brown and form sheaths

that wrap completely around the branchlet. Tiny flowers are on long spikes. The fruits look like grapes. Distribution: Along both coasts throughout, typically growing only on ocean beaches, outside the forest, often in association with the mangrove Conocarpus erectus (Combretaceae). Recognition: In its limited habitat, this species is unmistakable. Coccoloba has 19 species in Panama, so this is another big genus to which we do not do justice. Leaves are highly variable (not within, but between species): narrow or wide, pointed or round, shiny or hairy, but the sheath is reliable. They are also highly varied in forest type, because there are species of dry areas, wet areas, and lower mountains.

Ruprechtia costata

Ruprechtia costata

Ruprechtia costata (rascador, dardo). A medium-sized tree of dry regions. The bark is gray and fissured. Twigs have lenticels. Leaves are simple, alternate, thick and stiff, rather bunched and irregularly placed around branches; margins are wavy and sometimes have a few teeth. Veins on the leaf under-

side are yellow and elevated. The sheath at the base of each leaf is small, and when it falls away not much of a scar is left. Seeds have three wings, and turn from yellow to reddish brown; when present they cover the tree. Distribution: Grows in open farmland of the Pacific slope in the driest zone of c. Panama and nw. Costa Rica; common at Sarigua National Park in Herrerra. Also appears sparsely on limestone outcrops around the Canal. Recognition: The inconspicuous leaf sheath makes this tree more difficult to pick out than other Polygonaceae. Without that sheath or fruits, Ruprechtia is not especially distinct, though not many species grow in its habitat. The distinctive winged seeds are just like those of Triplaris, though darker when mature.

Polygonaceae

Coccoloba uvifera

Ruprechtia costata

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Proteaceae

Triplaris cumingiana

Triplaris cumingiana

Triplaris cumingiana (palo santo, guayabo hormiguero, vara santa). A medium-sized tree of forest edge. The trunk is straight, usually with small buttresses and somewhat fluted, and the bark peels away to leave large irregular patches of light-colored inner bark. Leaves are simple, alternate, spaced regularly along branches, and have prominent straight and parallel secondary veins. Should you break off a branch, you will discover that the inside is hollow and likely inhabited by ants that have a nasty sting— you will not do it again. Tapping stems or branches on saplings may get the ants out. The flowers are small and white or greenish, produced on long stalks above the crown. After they fall, the seed left behind grows three long, red wings. Because the red seeds

remain on the tree for awhile, they are conspicuous from a distance and look like bright red flowers. Distribution: Widespread in lowlands near the Canal, especially toward the Pacific, and seen in other dry areas near the Pacific coast. Not known in Costa Rica. Often an edge species, and there are many along roads and open areas of the Pacific side of the Canal Area, where the bright red seeds are easy to spot toward the end of the dry season. Also occurs occasionally in mature forest, but saplings are not seen in the shady understory. Recognition: A good species to learn well: watch out for saplings and their ants! The leaves look a lot like those of Annonaceae, but the leaf sheath can usually be found and is nothing like Annonaceae. The straight secondary veins of Triplaris are quite different from those of Coccoloba, and Ruprechtia has much smaller leaves. The fluted, peeling trunks on big trees are easy to spot, but watch for Quararibea (Malvaceae—Bombacoideae). The three-winged seeds are easy to spot on the forest floor when falling. A second Triplaris species, T. americana, is rare and known only in far e. Panama.

Roupala montana

Proteaceae Roupala montana

A large family worldwide, with 1700 species, but the vast majority are in heathlands of Australia and South Africa, where they are the dominant plant family. It is the family of the Australia macadamia nut in the genus Macadamia. In tropical forest, they are unimportant and seldom common. There are 86 species in tropical America and 7 in Panama, of which we cover just 1. The family is best characterized by its flowers, which resemble bottlebrushes (but are not the bottlebrush trees, which are Myrtaceae). Many have compound leaves, but many do not, and no simple character apart from flowers distinguishes the family.

Roupala montana (carne asada, árbol carne, ratón). A medium-sized tree of forest and savanna. The trunk can be straight and cylindrical, and large trees have small buttresses, but trees growing in open areas often have twisted trunks that branch near the ground. The bark is light brown, with dark brown lenticels. Leaves are alternate and rather

thick, but come in two very different forms—some are compound and some are simple, and a single branch can have both types. The simple leaves are heart-shaped, with a long, pointed tip, sometimes but not always toothed, and rather variable in shape. In compound leaves, there are 5–12 leaflets, with 1 terminal. The leaflets are toothed and can be opposite or alternate along the rachis, and are also rather variable in shape. Crushed leaves, especially young ones, smell like hamburger or beef, hence the common names. Flowers are yellow, arranged around a spike like a bottlebrush. Fruits are dry capsules carrying winged seeds. Distribution: Mainly a species of dry areas of the Pacific slope, where it can occur in forest or savanna, but also shows up in lower mountains. It can be found, though rare, in moist forest at Barro Colorado I. and near Gamboa. Recognition: The leaves are distinctive once they have been seen a few times, being rather thick and at least some having conspicuous teeth. When compound the coarse and irregular teeth are distinctive. The meat smell is subtle, and not everyone gets it. Dendropanax arboreus (Araliaceae) also has heart-shaped leaves that are sometimes lobed or toothed, rather similar to the simple leaves of Roupala. Two other Roupala species are found in Panama. One is rare and known only in e. Panama, and we have seen it in the upper Chagres region east of Panama City. The other is in mountains of w. Panama and Costa Rica, generally at higher elevation than R.

Proteaceae

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Triplaris cumingiana

Roupala montana

montana (ironic given the species name). All are known for their meat odor and have similar leaves. The other four species of Proteaceae in Panama are in the genus Panopsis, and their leaves do not resemble those of Roupala. They are simple, opposite or subopposite, without teeth, and are shiny and

glossy green; the tertiary venation is finely reticulate. They are montane and cloud-forest species. P. suaveolens is around Cerro Campana and Cerro Azul near Panama City, and can be found widely but sparsely in mountains through Panama and Costa Rica. The remaining species are seldom collected.

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Putranjivaceae Drypetes standleyi

Putranjivaceae

(Formerly Euphorbiaceae)

Drypetes standleyi

A small family of about 200 species, mostly in the Old World, with just 15 in the New World. Formerly, they were considered part of the Euphorbiaceae. There are three species of trees in a single genus in Panama. With so few species, we do not consider family traits.

Drypetes standleyi (madroño). A tall tree of moist and wet forest. The trunk is straight and cylindrical, and bark is gray with horizontal scars and sometimes white pores (lenticels). Leaves are simple, alternate, small, and glossy green above. At least some leaves might show slight asymmetry, with left (or right) side narrower or more curved than the oth-

er. The leaf underside is lighter in color than above, and there are only a few secondary veins arching forward. There are often small brown cancer-like growths on the leaves (insect galls). Flowers are small, light green, in groups on short stalks at leaf axils. Fruits are fleshy and turn from green to yellow or brown. Distribution: In lowland wet and moist forest, especially in Costa Rica, though fairly common in the Canal Area. Recognition: Might qualify as the least recognizable tree in Panama. Lacks all distinctive features: no hairs, no latex, no odor, etc. The slight leaf asymmetry and bark scars are weak characters, but might help given the otherwise poor distinctions. Around Barro Colorado I., the little insect galls are useful, but not elsewhere. Might be confused with Annonaceae (see Oxandra), Chrysobalanaceae (see Licania), Lacistemataceae (see Lacistema), Lauraceae (see Ocotea cernua), Salicaceae (see Laetia, although it has teeth), Pera (Euphorbiaceae), or Margaritaria (Phyllanthaceae). Two other Drypetes species are known in Panama, one known only in the far west and in Costa Rica, the other only along the Caribbean coast of e. Panama.

Rhamnaceae Colubrina glandulosa

Colubrina glandulosa

A moderate-sized family, with about 900 species, most of which are trees or shrubs, though there are some lianas and a few herbs. Some of the species are in temperate zones, but most are tropical. There are far more in the Old World, with only 170 in the American tropics. North America and Europe have the buckthorn genus, Rhamnus, and California has the mountain-lilac genus, Ceanothus. Ten species of trees and shrubs are known in Panama, and we cover 4. The rainforest species of Rhamnaceae are not easy to characterize. Some have alternate leaves, but some have opposite, and though the family is known for spines, two of the species we treat are not spiny. Many Rhamnaceae have three major veins arising from the leaf base, but only two of the three species we cover share this trait. The flowers are generally recognizable, having five small petals that curve inward into a claw, with a stamen attached to the petal (as in Ceanothus). Without flowers, the tropical trees have to be learned on their individual characters. Be-

cause Panama’s species are uncommon and seldom seen, the family is difficult to learn.

Colubrina glandulosa (carbonero, frio). A medium-sized tree of open areas. The trunk is straight, usually cylindrical but sometimes a bit fluted. Juveniles have long, thin branches. Leaves are opposite or subopposite (i.e., not quite opposite), dark green above and light green below, with veins yellow and prominent below; two or four of the secondary veins are large and originate at the leaf base. There is a pair of small, green, rounded glands where the leaf meets the petiole. Flowers are small, greenish yellow, in tight groups at leaf axils. Fruits are three-parted capsules that turn from green to brown then split open. Distribution: Principally in open areas and small woodlots and farmland of rural Panama, particularly in the Canal Area but also to the west, with a few records in Costa Rica. Very rare in forest. Recognition: The opposite leaves and prominent secondary veins suggest Pittoniotis trichantha or Coutarea hexandra (Rubiaceae), but the bulbous glands and lack of stipules between the leaf pairs distinguish Colubrina from any Rubiaceae. There is a liana with very similar leaves, Omphalea diandra in the Euphorbiaceae, but it should only be confused in the juvenile state. Colubrina has heavy wood prized in rural communities, where the tree is common. Because it grows rapidly in farmland, it appears to be an excellent candidate for plantation forestry.

Rhamnaceae

Drypetes standleyi

Colubrina glandulosa

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Rhamnaceae

Colubrina heteroneura

Colubrina heteroneura

Colubrina heteroneura (espino del diablo). A

small spiny tree of dry areas. The trunk is sometimes irregular, and bark peels off in vertical strips. Branchlets have spines and fine reddish hairs. Leaves are dark and shiny, alternate and regularly spaced along branchlets. The base of each leaf, adjacent to the pet-

iole, has a little gland on the margin. Flowers and fruits are like those of C. glandulosa. Distribution: Nearly exclusively around the Canal Area, and fairly common in open areas or secondary forest around Panama City and near Madden Dam. Recognition: Not at all like C. glandulosa, which has opposite leaves and no spines. There is a Rubiaceae with similar spines, Chomelia spinosa (not illustrated), but with opposite leaves. Other small spiny trees in Panama have much different leaves. Two more Colubrina species are known in Panama. C. spinosa has leaves with glands much like those of C. glandulosa and is widely collected in Caribbean Costa Rica but barely into Panama. The other species is very rarely seen.

Rhamnus sharpii

Rhamnus sharpii

Rhamnus sharpii. A medium-sized tree of high

mountain forest. The trunk is straight and cylindrical, with light-colored bark. Branches are finely pubescent. Leaves are simple, alternate, sometimes slightly toothed but often not, light blue below. Secondary

veins are straight and closely parallel, yellowish, and raised on the underside. Flowers are small, greenish, in axillary clusters. Fruits are dark berries. Distribution: High mountains of far w. Panama into Costa Rica, where it can be common. Recognition: The bluish leaves with prominent yellow veins are easy to spot, but remarkably similar to those of Alnus acuminata (Betulaceae) in the same habitat. Alnus is a big tree, however, with woody fruits, and the leaves are much more conspicuously toothed. Two more Rhamnus species are found in the mountains of w. Panama and Costa Rica. Neither has bluish leaf undersides, and they resemble the north temperate buckthorns.

Rhamnaceae

Colubrina heteroneura

Rhamnus sharpii

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Rhizophoraceae

Zizyphus mauritiana (guindo). A small spiny

tree from Asia. Branchlets have a pair of spines at the base of each leaf, one curved, one straight. Leaves are small, round, alternate, whitish beneath, with three main veins arising together at the leaf base. The fruit is round, plum-sized, and matures yellow. Distribution: Native to Asia, but grows in rural area all over the tropics, cultivated for its sweet fruit. Not in forest. Recognition: The leaves and spines are easy to recognize, very similar to those of some Californian Ceanothus. One species of Lauraceae, Cinnamomum triplinerve, has three main leaf veins, but its leaves are bigger and it has no spines. The genus Strychnos (Loganiaceae, a

family not covered in the main text) has similar leaves, but opposite. Zizyphus has two native species in Panama, both uncommon and both with leaves having the same distinctive set of three main veins. Z. chloroxylon is found at the Santa Rita ridge east of the Canal and on the Osa Peninsula in Costa Rica, but nowhere else. It is a huge canopy tree, with shiny green leaves. The other species is known only in the upper Chagres region and the Darien. One more genus in the Rhamnaceae, Rhamnidium, has just one record from Bocas del Toro in Panama.

Rhizophoraceae Cassipourea elliptica

Cassipourea elliptica

A small family of tropical trees and shrubs, with just 130 species worldwide and only 18 in tropical America. It is the major mangrove family in the New World, though other genera in the family grow in upland forest. There are just three species in Panama, and we cover two. Rhizophoraceae in general have opposite leaves and might be confused with Myrtaceae. Beyond this, the two Panama genera are not obviously similar and are best learned by their individual traits.

Cassipourea elliptica (huesito, limoncillo, ajo). A medium-sized forest tree. The trunk is straight, cylindrical, and the bark has white lenticels. Branches are opposite one another along the trunk, and branchlets are swollen at the leaf nodes, showing clearly where leaves have fallen. Leaves are opposite and simple, often toothed, though sparsely and sometimes not at all. Each secondary vein arches forward to meet the next secondary vein, making a collecting

vein near the leaf margin that is evident on the leaf underside. Flowers are white, with fringed petals, in small clusters at leaf axils. Fruits are capsules that turn yellow as they mature, splitting open to reveal seeds covered in a white arils. Distribution: A very widespread species, in all types of forest from moist and wet to lower montane, but not in the driest zone. In swampy forests it can be one of the dominant species, along with Prioria copaifera (Fabaceae—Caesalpinioideae). Not seen along roads or in open areas. Recognition: Toothed, opposite leaves are distinctive, but unfortunately the teeth on this species are sometimes inconspicuous. If you look carefully at many leaf margins, usually at least some will appear toothed. Another species with these traits is the very similar Crossopetalum parviflorum (Celastraceae), which also has opposite leaves that only sometimes have teeth. A good detail to separate the two is branchlet form: slightly grooved in Cassipourea but not in Crossopetalum. See also Siparuna pauciflora (Siparunaceae) and Mollinedia (Monimiaceae, mentioned under Siparunaceae), both have opposite leaves with teeth; Siparuna has a strong and unusual odor but Mollinedia does not. If no toothed leaves are found, opposite leaves without latex might pass for Myrtaceae and Eugenia coloradoensis especially might be confused with Cassipourea. The way branches are swollen where they meet the leaves in Cassipourea will be very helpful if you remember to check, but it is a subtle trait.

Rhizophoraceae

Zizyphus mauritiana

Cassipourea elliptica

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Rubiaceae

Rhizophora mangle

Rhizophora mangle

Rhizophora mangle (mangle rojo, mangle colo-

rado, red mangrove). America’s dominant mangrove. A tall tree of salt marshes, with a straight, cylindrical trunk and spectacular stilt-roots. In large individuals, the roots can be more than 2 m long and branched; in well-developed forests the stilts form a continuous jungle gym at the base of the trees (and traversing a mangrove stand can only be accomplished by climbing the stilts). The bark is smooth and reddish brown. Leaves are simple, opposite, densely clustered toward the branch ends, shiny green, with secondary

veins barely visible; they often have small black spots. The seed germinates and grows a small stem while the fruit is still on the tree, so that fruits have long greenish spikes. The seed floats, but with one heavy end down, until it roots in mud. Distribution: This is the mangrove closest to the sea’s edge, and extensive stands are widespread on both Caribbean and Pacific coasts. It nearly always grows in pure stands, with no other trees mixed in, and never grows away from the sea. Recognition: Rhizophora is unmistakable through the Americas: the other mangroves lack stilts. Small individuals lacking stilts are seldom seen; their leaves are distinctive in being densely clustered at the branch tips. Mangroves have very hard wood frequently used in construction and for making charcoal. Rhizophora includes two more species, but it turns out one is a hybrid of the others. They are differentiated only on the basis of flower size and number; both are much less common than R. mangle.

Rubiaceae Alibertia edulis

Alibertia edulis

This is the fourth largest plant family in the world, with 13,000 species, ranking after the Asteraceae (daisies and sunflowers), the Orchidaceae, and the Fabaceae (legumes). It is the coffee family, and coffee (genus Coffea) is widely planted in Panama and Costa Rica but is native to Africa. Most species in the family are small trees or shrubs of tropical forest understory, and because many are very common, it is not difficult to make the case that this is the most important tropical forest family: nearly everywhere in the world’s tropics, the forest understory has many Rubiaceae. Indeed, because most Rubiaceae are shrubs or trees, whereas many Fabaceae are herbs, Rubiaceae matches Fabaceae in species richness among tropical tree families. In Panama, there are 236 species, compared to 234 Fabaceae and 143 Melastomataceae. Thus, Rubiaceae is 10% of Panama’s tree checklist. We cover 32, with 30 illustrated. Because the family is abundant and can always be recognized from one subtle trait, we suggest that it is the most important tropical forest family to learn (not counting the really easy-to-spot palms and melastomes). All Rubiaceae have opposite leaves without teeth, and all have a tiny but crucial character called an interpetiolar stipule. The stipule, recall, is the tiny leaflike appendage at the base of

each leaf that many plants have. In Rubiaceae, this stipule stretches across the branch between the pair of petioles. This is very distinctive when the stipule remains. Moreover, the form of the stipule is highly variable across Rubiaceae, sometimes a simple triangle and sometimes forked or lobed, and can be a useful trait in identifying genera or species. To best study the stipules, look at the tip of growing branches, where new ones have just appeared. Even after stipules fall off, the distinctiveness holds because there is a scar stretching across the branch between the petioles, and because stipules often fall, this scar is the real character you rely on to identify Rubiaceae. The only glitch, and it is small, is that a few non-Rubiaceae show interpetiolar scars, most notably Achanthaceae and Lamiaceae, plus the genus Cassipourea in the Rhizophoraceae. Because many Rubiaceae are common in forest understory, you can quickly impress friends by pointing out any opposite-leaved shrub or treelet in a tropical rainforest (or even botanical garden) that has interpetiolar stipules and identify it as Rubiaceae. Flowers are tubular, sometimes very long, and usually white, though there are species with yellow, orange, and even blue flowers. Fruits come in two types: fleshy berries or capsules holding winged seeds.

Alibertia edulis (trompito, madroño, zumbo,

guayabito de monte). A treelet of the dry zone. The small trunk typically branches near the base. Leaves are simple, opposite, pointed at the tip but rounded at the base. Stipules are pointed and persistent, starting green but remaining after they turn brown; they form a tube around the tip of the growing branchlet. There are sometimes tiny pits (called domatia) visible on the underside of the leaf, at the axils between

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Rhizophora mangle

Alibertia edulis

the primary and secondary veins. Flowers are fairly large, white, tubular, with four petals. Distribution: Mostly on the Pacific slope, where it occurs in open shrublands and secondary forest. Uncommon in old growth at Barro Colorado I. and Soberania, and occurs less commonly in similar habitat on the Caribbean slope, but not inside wet forest. Recognition: Alibertia has nondescript leaves and is not easy to learn. The persistent stipules are helpful (many Rubi-

aceae lose their stipules before they are brown), but you have to study leaf and stipule shape carefully. Several Psychotria have similar leaves with domatia, and the rare Borojoa panamensis (not illustrated) has very similar leaves. Two other Alibertia species are found in Panama, one primarily in montane forest, the other very rare.

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Alseis blackiana

Alseis blackiana

Alseis blackiana (mameicillo). A medium-sized

forest tree. The trunk of larger trees is straight but slightly fluted, with small buttresses. Bark is light brown, with fine fissures, soft (you can sink a finger into it slightly). Leaves are opposite, long and fairly narrow, widest beyond the middle, with a very short petiole, and have prominent yellow secondary veins. Stipules fall off quickly, leaving a scar around the branch. As juveniles, there is a single vertical stem with each leaf perpendicular to it; successive pairs of leaves are perpendicular to one another. In the shade, these successive pairs can be extremely close together, so four leaves can appear to arise from the same spot on the stem. Flowers are white or yellow-

ish, small and inconspicuous in the canopy. Fruits are unusual, in clusters of tiny, narrow brown pods. They look like a dried branchlet from a spruce tree (the northern conifer used for Christmas trees)— each little seed pod looks like a spruce needle. Distribution: Seen widely in the Canal Area and abundant in old forest at Soberania and Barro Colorado I. Otherwise, known from only a few collections. Saplings are numerous in the shaded forest understory. Recognition: This is one of the few Rubiaceae that grows to real tree size. The long, opposite leaves, broader beyond the middle, are distinctive, and easy to spot in the canopy; coupled with the soft bark, adults can thus be learned pretty easily. The nearly whorled arrangement of leaves on stems of juveniles is also easy to recognize. A rather similar Rubiaceae is Genipa americana, but it is scarce inside the forest and has much different bark. Alseis is probably more often confused with Anacardium excelsum (Anacardiaceae), Gustavia superba (Lecythidaceae), or Pouteria fossicola (Sapotaceae). All have long leaves wider above the base, but alternate not opposite. Alseis may be confusing when the leaves are very crowded and it is not obvious they are opposite.

Amaioua corymbosa

Amaioua corymbosa

Amaioua corymbosa (madroño). A medium-

sized tree often of secondary forest. The trunk is typically highly fluted. Bark is brown or reddish, peeling in small strips. Leaves are opposite, round in shape, with prominent yellow veins below. There can be little domatia along the main vein underneath. The

stipule is brown with fine hairs; the scar it leaves when falling has fine teeth along the top. White flowers are tubular. Distribution: Mostly on the Pacific slope, and very common in secondary forest in the Pacific half of the Canal Area; also in old growth there, but less common. Rarely known away from the Canal. Recognition: The small, fluted trunk looks like that of Macrocnemum, which has similar leaves but much different stipules. The best character for Amaioua is the tiny, brown, toothed stipule scar. Two other Amaioua species are found in Panama, one restricted to wet forests of the Canal Area, the other to lower mountains from c. Panama through Costa Rica. They have similar leaves, though in one larger, the other smaller, than in A. corymbosa.

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Alseis blackiana

Amaioua corymbosa

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Arachnothryx buddleioides not found

Arachnothryx buddleioides

Arachnothryx buddleioides (dos caras). A

cloud-forest treelet. Leaves are simple, opposite, conspicuously brown on the underside with prominent, raised veins. Flowers are very small, pinkish, on long, terminal spikes. Fruits are tiny, dry capsules that split

open. Distribution: Cloud forests from c. Panama through Costa Rica. Recognition: Most easily confused with its namesake, Buddleja americana (Scrophulariaceae), which has Rubiaceae-like stipules and brown leaf undersides, though less conspicuous venation. Otherwise, Arachnothryx buddleioides might only be confused with some unusual Myrtaceae with brown leaf undersides, but of course those lack the Rubiaceae interpetiolar stipules. Three more Arachnothryx species are found in Panama, mostly in montane and wet forest. Some have silvery, not brown, leaf undersides, but others have typical green leaves.

Calycophyllum candidissimum

Calycophyllum candidissimum

Calycophyllum candidissimum (madroño, ala-

zano, lluvia de plata, salamo). A medium-sized tree of the dry zone. The trunk is straight but fluted, and bark is gray or reddish brown, with vertical strips peeling off to reveal smooth, reddish inner bark. Leaves are simple, opposite, rounded and nearly as wide as long but pointed at the tip, with widely spaced secondary veins. The stipules are conical and cup-shaped, but quickly fall off, leaving a conspicuous scar behind that rings the branch at the base of each leaf pair. The flowers are small, white, in clus-

ters above the leaves; beneath the flowers are bright white calyx lobes that make the whole display visible from a distance. Fruits are capsules that split to release winged seeds. Distribution: This is one of the characteristic species of drier regions, common and conspicuous in farmland and along roads near the Pacific, and abundant along the Canal from Panama City to Gamboa. It typically associates with Bursera simaruba (Burseraceae), and both species also occur on limestone outcrops near the Caribbean. Recognition: Best known from the bright white floral lobes that are very conspicuous at the start of the dry season. The rounded leaves and red trunk together are good characters. Pittoniotis trichantha has rather similar leaves, but not peeling bark; Alibertia edulis has peeling bark but different leaves. Calycophyllum might be confused with Terminalia oblonga in the Combretaceae, which has smooth bark and vaguely reminiscent leaves. When flowering, Calycophyllum can be easily seen at great distances.

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Arachnothryx buddleioides

Calycophyllum candidissimum

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Cosmibuena macrocarpa

Cosmibuena macrocarpa

Cosmibuena macrocarpa (tabaquillo). A small

cloud-forest tree. Leaves are opposite, thick, smooth, with barely visible secondary veins. Flowers are very long, cream-colored tubes. Distribution: Wet and lower montane areas of c. Panama, and also grows near sea level on the Caribbean coast. Common at

Cerro Campana. Seldom seen in Costa Rica. Fruits are dry capsules; seeds are tiny and winged. Recognition: The leaves are highly distinctive and can only be confused with those of the genus Clusia in the Clusiaceae, which is remarkably similar. In Clusia, milky latex emerges slowly from a broken leaf, but Cosmibuena has no latex (nor does any Rubiaceae). Two other Cosmibuena species are found in Panama. One has thick, narrow leaves with inconspicuous veins, and white flowers with pink tips. The other has thick leaves, but with prominent secondary veins. Both are largely montane species, but one reaches lowland wet forests.

Coussarea curvigemmia

Coussarea curvigemmia

Coussarea curvigemmia (huesito). A forest treelet. Leaves are opposite and have a distinct shape at the base. Just before the leaf narrows to a point, it ex-

tends toward and almost touches the branchlet. There are tiny domatia at the vein axils on the leaf underside. Flowers are white tubes; fruits are berries that turn white when mature. Distribution: Mostly known in the Canal Area, and abundant in old growth at Barro Colorado I. Elsewhere seldom collected. Recognition: The leaf base is very distinct. Another 14 other species of Coussarea are known in Panama, but many are very rare. Several are from montane and wet forests, and some have narrow ranges. Some of the species, but not all, share the extended leaf base.

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Cosmibuena macrocarpa

Coussarea curvigemmia

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Coutarea hexandra

Coutarea hexandra

Coutarea hexandra (azulejo, quina). A small forest tree of the Pacific slope. Branchlets have conspic-

uous white lenticels. Leaves are opposite, wide and rounded, with prominent parallel secondary veins. Stipules are small, brown triangles. Flowers are white with pink lines. Fruits are dry capsules that split to release winged seeds. Distribution: Especially on the Pacific slope of c. Panama and nw. Costa Rica. Seen throughout the Canal Area, but not common. Recognition: This species is uncommon and not easy to learn. Wide leaves with clear secondary veins are much like those of Exostema mexicanum and Pittoniotis trichantha, and Colubrina glandulosa (Rhamnaceae) likewise has similar leaves.

Exostema mexicanum

Exostema mexicanum

Exostema mexicanum (azulejo, quina). A medium-sized forest tree. Branchlets have white lenticels.

Leaves are opposite, broad and rounded, and have conspicuous, parallel secondary veins. There are tiny domatia in the vein axils of the leaf underside. Stipules are small triangles and remain on the branch even when old. Flowers are white; the fruits are dry capsules with white lenticels. Distribution: At drier sites, especially in nw. Costa Rica. In Panama, found on limestone near the Madden Dam and a few other sites. Recognition: See Coutarea hexandra and other species mentioned there; Exostema and Coutarea are very similar, though Exostema is a larger tree.

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Coutarea hexandra

Exostema mexicanum

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Faramea luteovirens

Faramea luteovirens

Faramea luteovirens (benjamín, huesito, garro-

tillo, jazmín). A wet-forest shrub or treelet. Leaves are opposite, thick and shiny, light green on the under-

side with a few widely spaced secondary veins. Stipules are tiny triangles, narrowing to an extended point; the pair growing at the tip of the branch, between the outer pair of leaves, has extended points that cross. Flowers are white, with short tubes and four petals. Fruits are blue berries. Distribution: Widespread in e. Panama’s wet forests, though not common. In the understory of mature forest. Recognition: The crossed stipules are a great character, but many Faramea have them. F. luteovirens has unusually thick leaves for Rubiaceae, like the very different Cosmibuena.

Faramea occidentalis

Faramea occidentalis

Faramea occidentalis (benjamín, huesito, garro-

tillo, jazmín). An abundant treelet of forest understory. The trunk is straight, branches are opposite and often found on the lower half of the trunk. Leaves are

opposite. The stipules have extended, filamentous tips, crossing in the terminal pair, like a tiny pair of scissors. Flowers and fruits as in F. luteovirens. Distribution: Widespread, especially in forests on the Pacific half of the isthmus, though also at lower montane sites and less commonly on the Caribbean slope. Always inside mature forest. It is the dominant treelet of the old-growth forest understory at Barro Colorado I., and common from there to Panama City. Recognition: Easy to learn from the crossed terminal stipules, though many Faramea share this trait. The other Faramea are wet-forest species, usually not overlapping with F. occidentalis.

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Faramea luteovirens

Faramea occidentalis

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Faramea papirifolia

Faramea papirifolia

Faramea papirifolia (cabeza de víbora). A wetforest treelet. Leaves are opposite, with many parallel secondary veins that are deeply impressed above and raised below; even the tertiary veins are raised below. Stipules are persistent, long triangles, rather thick and turning brown when older; they do not cross. Distribution: Known only in wet areas and low mountains just east of the Canal. Recognition:

The deeply impressed veins are conspicous and easy to learn. Stipules do not cross as in other Faramea. Another Faramea, F. papillata, is also confined to the Cerro Jefe area, and has raised secondary veins, but its tertiary veins are smooth. A third Faramea is known only near Cerro Jefe, and it takes the name F. jefensis, but it does not have impressed veins. One widespread Faramea of wet forests, F. suerrensis, has impressed veins, but it also has two conspicuous marginal collecting veins and looks remarkably like Melastomataceae. The genus Faramea has 15 more species in Panama; three of these were listed under F. papirifolia. Most are montane or wet-forest species, but many are seldom seen. Stipules are pointed but not always crossed. Flowers are similar across the genus, but in several species are bluish.

Genipa americana

Genipa americana

Genipa americana (jagua). A medium-sized tree of open areas. It often branches near the ground, and smaller branches have clear scars marking the location of fallen leaves. Leaves are opposite, fairly large, broadest beyond the middle, and tend to be clustered toward the ends of the branches. Stipules are broad, triangular, brown, often persistent. Flowers

are short yellow tubes. Fruits are large, round, fleshy, edible. Distribution: Essentially a species of disturbed areas and farmland, very widespread but mostly on the Pacific slope. Rare inside the forest. Recognition: This is one of the few Rubiaceae that grows to real tree size. Inside the forest, see Alseis blackiana, which has similar though longer and narrower leaves. Tocoyena pittieri (not illustrated) also grows in open areas and has similar leaves; Tocoyena has very long, tubular flowers, and leaves are smooth and glossy green, unlike Genipa. Fruits of Genipa are used for a purple dye. You may run across two other Genipa species names in Panama, but they were recently reclassified into the genus Agouticarpa.

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Faramea papirifolia

Genipa americana

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Guettarda crispiflora

Guettarda crispiflora

Guettarda crispiflora (salvia de montaña). A

small tree of cloud forest. The bark is light colored and peels off in small pieces. Leaves are opposite,

broad and sometimes almost heart-shaped, with a long pointed tip; the underside is light green and has conspicuous, yellow, arching secondary veins. The stipule usually falls off. Leaf, petiole, and stipule all have tiny, fine white hairs. Flowers are white tubes with fringed petals. Fruits are largish berries, green turning purple. Distribution: Mostly in lower montane forests, most common in Costa Rica. In Panama known from Cerro Campana and Chiriqui. Occasionally found at wet, lowland sites. Recognition: The fuzzy leaves are easy to spot. Pittoniotis trichantha has very similar leaves, but occurs only in the lowlands.

Guettarda foliacea

Guettarda foliacea

Guettarda foliacea (guayabo de monte, espino

amarillo). A medium-sized tree of moist to wet forest. Small trunks and young branches sometimes have opposite spines, but not larger trees. Twigs are lightly pubescent. Leaves are opposite, clustered toward

branch ends, and usually bullate, with veins impressed above. Stipules are long, narrow, and usually remain after turning brown. Flowers are white tubes, and fruits are large berries. Distributed: Widespread in moist to wet and lower montane forest throughout Panama and Costa Rica. Common in old-growth and secondary forest around Barro Colorado I. and Soberania. Recognition: When lacking spines, a fairly nondescript Rubiaceae. The impressed veins are useful. Look for the long, narrow stipules. Five more Guettarda species are found in Panama, including species from montane, dry, and wet forests. All are treelets.

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Guettarda crispiflora

Guettarda foliacea

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Hamelia patens

Hamelia patens

Hamelia patens (guayabo negro, canelito). A

weedy treelet or shrub. Leaves come in groups of three or four. Flowers are orange tubes and nearly always present. Distribution: Very widespread, dry to

wet and montane, along roads and forest edge. Recognition: Usually known by its flowers, which can be seen throughout the year. The whorled leaves (meaning more than two emerging from the same spot) are unusual in any family. The shrub that might be confused with H. patens is Isertia haenkeana (not illustrated), an abundant roadside weed; it has larger leaves with clearly raised secondary veins. See also Rauvolfia littoralis (Apocynaceae) with whorled leaves but latex. Five more Hamelia species are found in Panama. A common one throughout the area, H. axillaris, has yellow flowers and leaves in pairs. The rest are rare.

Ladenbergia macrocarpa

Ladenbergia macrocarpa

Ladenbergia macrocarpa (buchirago, michura-

go, fruta de mono). A medium-sized tree of cloud forest. Larger trees have small buttresses and black, fissured bark. Leaves are opposite, large, rounded, light colored, with reddish veins on the underside. Flow-

ers are large, cream-colored. Fruits are dry capsules holding winged seeds. Distribution: Mountains of c. and e. Panama. Not known in Costa Rica. Recognition: The round leaves with reddish veins are distinctive in the Rubiaceae. Leaves are reminiscent of those of Coccoloba caracasana (not illustrated) and C. uvifera (Polygonaceae), but those have alternate leaves. Five more Ladenbergia species are found in Panama, and several—but not all—have large, rounded leaves. Two are montane and seen widely in Costa Rica; the rest are very rare and found only at a few sites in Panama.

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Hamelia patens

Ladenbergia macrocarpa

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Macrocnemum roseum

Macrocnemum roseum

Macrocnemum roseum (palo cuadrado, madroño, canaleto). A medium-sized forest tree. The trunk is straight but highly fluted. Leaves are fairly large, opposite, wide and rounded toward the tip and narrow at the base; the petiole is very short so the base of the leaf almost touches the branch. The stipules are conspicuous at the apex of the branch, between the final pair of leaves: large, oval, and green. But they fall quickly, leaving a scar between leaf pairs further down the branch. Flowers are small, bright

pink, in dense clusters. Fruits are dry capsules holding very small, winged seeds. Distribution: Widespread in moist and wet lowland and some lower montane sites. Common in the Canal Area. Found in mature forest, though not common, and often along wooded roadsides and rivers. There are a number of trees along Pipeline Rd. that drop bright pink flowers on the road. Recognition: Inside the forest, quickly spotted from the unusual trunk. Only two other species have such a trunk, Amaioua corymbosa (also Rubiaceae) and the big Fabaceae (Papilionoideae) Platypodium elegans. The latter has compound leaves with tiny leaflets and is almost never seen as a small tree. Amaioua might be confused, but it has rounder leaves and typically a less fenestrated trunk than Macrocnemum. In juveniles with small stems, Macrocnemum could be confused with several other large-leaved Rubiaceae, such as Genipa americana or Posoqueria latifolia, the latter especially in having a large stipule.

Palicourea guianensis

Palicourea guianensis

Palicourea guianensis (recadito). An abundant treelet of forest gaps. Leaves are opposite, fairly wide, though not as wide as long. Stipules are persistent, green, and forked, each consists of two long triangles; there thus appear to be four separate stipules at each leaf pair. Flowers are yellow, tubular, on orange stalks. Fruits are small berries. Distribution: Common and widespread in lowlands, except the driest zone. Abundant around the Canal. Always in small

natural clearings where there is lots of light. Recognition: The most common Rubiaceae in gaps in lowland forest; usually the fairly wide leaves identify it. Most Psychotria have narrower leaves and white flowers. See Hamelia axillaris (not illustrated, but see H. patens), another forest shrub with similar leaves. The forked stipules are a very good character; all Palicourea have them. Isertia (not illustrated) has similar stipules, though divided to the base, and some Psychotria have forked stipules. Palicourea is a very big genus, with many species at the margin of tree versus shrub status. We count 14 species as trees or treelets, and 18 more as smaller shrubs. They have orange, yellow, or even blue flowers, and all have forked stipules (though sometimes short). Most are montane, in fact Palicourea can be thought of as Psychotria of montane forest.

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Macrocnemum roseum

Palicourea guianensis

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Pentagonia macrophylla

Pentagonia macrophylla

Pentagonia macrophylla (teta de chola). A small forest tree. The trunk often has low forks and is sometimes leaning. Small branches are square in cross section. Leaves are opposite, very large, wide, and stiff. The stipules are triangular, and red when new; they are persistent, but turn brown when old. Flowers are small yellow tubes, sitting in dense clusters of red bracts at the ends of branches. Most trees have the bracts present, usually without flowers. Fruits are fleshy and turn red, held in the same dense

clusters at the ends of branches. Distribution: Common and widespread in lowland forests near the Canal, but rare elsewhere and barely reaches Costa Rica. Generally in forest shade, though commonly seen along Pipeline Rd. Recognition: Best recognized from the dense heads of red floral bracts that are nearly always present. The large, stiff leaves are also distinctive; some epiphytes in the family Araceae (Monstera and Philodendron) have rather similar leaves. Our checklist of Panama trees has nine Pentagonia species, but most are rare. P. gymnopoda is fairly common in wet forests near the Canal, and is remarkable among Rubiaceae in having leaves with such deep lobes that they look compound. As in Palicourea, there are additional species in the treelet– shrub gray area; a total of 15 species of Pentagonia are known in Panama.

Pittoniotis trichantha

Pittoniotis trichantha

Pittoniotis trichantha (candelo). Not illustrated. A medium-sized roadside tree. The trunk usually leans some and is often forked, and bark typically peels in small pieces. Leaves are rather wide, rounded at the base and thus can be nearly heart-shaped.

Secondary veins are prominent, white, and arch well forward. The stipule is a small, persistent triangle. Branchlets and stipules have soft white hairs, but leaves do not. Flowers are off-white, in dense clusters, abundant at the end of the dry season. Fruits are red berries. Distribution: Very common near Panama City but seldom seen elsewhere. Seen along all wooded roads on the Pacific half of the Canal Area. Recognition: Near Panama City, this tree can be learned well from the prominent veins on rather short, rounded leaves. Other Rubiaceae that reach this size (15–20 m tall) have longer, narrower leaves (Alseis, Genipa, Posoqueria, and Tocoyena, the latter not illustrated).

Pogonopus exsertus

Pogonopus exsertus

Pogonopus exsertus (sanguinaria, cresta de gallo). A treelet of moist to wet zones. Branchlets have white lenticels. Leaves are simple, opposite, broad

and round in the middle but narrowly pointed at both ends. Stipules are small triangles that usually fall off. Flowers are long pink tubes, each having a round calyx lobe adjacent that is bigger and darker than the flower. Fruits are capsules that split open. Distribution: Just a few records from the Canal east, and in Costa Rica. Fairly common on limestone near Madden Dam. Recognition: Usually known by the pink flowers with conspicuous round bracts. Warszewiczia coccinea also has bracts like this, though bright red, and it is a larger tree. Leaves of Pogonopus are like those of many Psychotria, but extending to an unusually narrow point at the base.

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Pentagonia macrophylla

Pogonopus exsertus

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Posoqueria latifolia

Posoqueria latifolia

Posoqueria latifolia (boca de vieja, borojó, fruta

de mono, tuliviejo). A medium-sized forest tree. Bark is gray, and in bigger trees it peels off in sheets. Leaves are opposite, thickened, dark shiny green above, and rounded at both tip and base. Stipules are narrow, moderately long triangles, and usually fall off. Flowers are very long, narrow, white tubes. Fruits are plum-sized and turn yellow; they are edible. Distribution: Very widespread, in all climates

from lowland to montane, in forests, farms, and even in cities; sometimes cultivated. Not especially common. Recognition: Usually known from the conspicuous flowers. Other Rubiaceae have white tubular flowers of the same shape, but none as long as those of Posoqueria. There is a species of Isertia, I. laevis (not illustrated), which might be confused: it has tubular white flowers nearly as long and is common along the outer part of Pipeline Rd., but has forked stipules. Another tree in the Rubiaceae, Tocoyena pittieri (not illustrated), has leaves like those of Posoqueria and bright yellow flowers of the same size and shape. Two other Posoqueria species are found in Panama, but one is extremely rare. Both have the same very long tubular flowers, rounded leaves, and long stipules.

Psychotria elata

Psychotria elata

shrublike branching is likely to be Psychotria. A rather large book could be devoted to the identification of the 106 species. We include just six of those, with five illustrated, to offer a brief summary of what the genus looks like.

Psychotria elata (sombrerito del diablo). A

Genus Psychotria. Nearly every tropical forest in the world has some Psychotria species in the understory, including Australia and even Hawaii. It is one of Panama’s huge genera, with 106 species, of which we include 54 in the tree checklist; the rest are shrubs. None are even small trees, though; all are treelets to shrubs, and deciding which of the 106 are trees was a matter of guesswork, based in some cases on only a few herbarium specimens. Nearly all are restricted to the shade of mature forest, and any small Rubiaceae in the shade with rather arching,

shrub or treelet of wet-forest understory. Leaves are opposite, shiny green, with veins raised below (impressed above). Stipules form a sheath enclosing the branch and are forked; they are often persistent. Flowers are small and white, but sit inside a large red bract. Fruits are oval berries that turn blue or black when mature. Distribution: Very widespread in lower montane and wet forests. The showy red floral display undoubtedly attracts botanists and leads to very thorough collecting. Recognition: Usually known by the red floral bracts, but it must be compared with P. poeppigiana (not illustrated), which also has red bracts.

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Posoqueria latifolia

Psychotria elata

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Psychotria grandis

Psychotria grandis

Psychotria grandis (cafecillo). A treelet of forest understory. The stem is green, straight and typically unbranched, with conspicuous ridges on four sides.

Leaves are fairly large, long, and have prominent yellow veins raised on the underside. Stipules are persistent and conspicuous. Flowers are white, very small, in big, loose, terminal clusters; fruits are red berries. Distribution: Widespread in lowland moist to wet forest. Common in forests throughout the Canal Area. Recognition: Very different from other Psychotria, which nearly all have shrublike, arching branches; P. grandis has a straight stem. The ribs on the stem are obvious and conspicuous. Leaves resemble those of Isertia haenkeana (not illustrated), but the latter is a roadside weed with orange flowers (see description under Hamelia patens).

Psychotria horizontalis

Psychotria horizontalis

Psychotria horizontalis (cafecillo). An abundant understory treelet or shrub. Branches are low and arching. Leaves are small, opposite, smooth and

shiny, and sometimes one member of each pair is a bit smaller than the other. Veins are slightly impressed above. Underneath the leaves, near vein axils, are tiny pits called domatia. Flowers are small and white. Fruits are red berries. Distribution: In lowland forests throughout, mostly on the Pacific slope and including dry regions. It is abundant in the understory of old-growth forest at Barro Colorado I. and Soberania. Recognition: This is the archetypal Psychotria, with smallish leaves, slightly impressed veins, and domatia. Close to the Canal, this is by far the most common Psychotria. In wet and montane forest, there are a number of other similar species.

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Psychotria grandis

Psychotria horizontalis

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Psychotria luxurians

Psychotria luxurians

Psychotria luxurians (cafecillo). A treelet of wetforest understory. Branchlets are nearly square in

cross section and yellowish. Leaves are simple, opposite, light green on the underside with conspicuous, raised, yellow secondary veins. Stipules are forked, persistent. Flowers are small and white; fruits are black berries. Distribution: Mostly a lower montane species, but also in lowland wet forests of the Caribbean slope. Recognition: Rather large as Psychotria go, with more of a treelet than shrub form. Conspicuous yellow veins and light-colored leaf undersides are good marks, but P. panamensis (not illustrated) is another montane species that is similar, though with unforked stipules.

Psychotria marginata

Psychotria marginata

Psychotria marginata (cafecillo). A widespread

understory treelet or shrub. Leaves are opposite, wid-

est near the tip, and narrow at the base. The leaf margin can look toothed with tiny filaments (cilia). Under the leaf there are tiny pits along the main vein. Flowers are yellow, fruits red. Distribution: In lowland forest of moist and wet zones. Common in most of the Canal Area, but less so than P. horizontalis. Recognition: P. marginata and P. horizontalis often grow near one another; P. marginata has larger leaves usually broadest above the middle. A much less common species, P. psychotriifolia (not illustrated), looks much like P. marginata, but has a conspicuous vein running along the leaf margin.

Rubiaceae

Psychotria luxurians

Psychotria marginata

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Rubiaceae

Psychotria poeppigiana

Psychotria poeppigiana

Psychotria poeppigiana (labios de puta, labios

ardientes, sombrerito del diablo, hot lips, mother-inlaw’s lips). Not illustrated. The simple, opposite leaves and branchlets are densely covered in soft pubescence. Flowers are small and yellow, held inside paired red bracts that are also very fuzzy and look

like lips, hence all the racy names. Distribution: Very widespread from moist through wet forest, reaching lower montane sites too. Abundant along Pipeline Rd. and in many secondary forests. Recognition: This and P. elata both have red bracts, but P. poeppigiana is fuzzy. They overlap broadly in distribution, but at the moist end P. poeppigiana is more common and at the montane end, P. elata. This is just a sampler, perhaps successful at revealing the range of variation in Psychotria. In fact, species of Psychotria are often distinctive and easy to learn. The difficulty is that there are so many, and without a lot of study you will leave many Psychotria unidentified.

Randia armata

Randia armata

Randia armata (rosetillo, jagua macho, mostren-

co, tres chucitos). A medium-sized, spiny tree. The trunk is typically irregular and branched near the ground. Bark is greenish or cream-colored. Branchlets have sharp spines that grow in opposite pairs and sometimes fork. Leaves are simple, opposite, often densely crowded at the end of branchlets, and have

an extended taper at the base. Flowers are white tubes. Fruits are golf-ball sized, with white lenticels, maturing yellow and fleshy. Distribution: Widespread though not common in moist to wet lowland and sometimes lower montane forest, but not in the very driest zone. Recognition: Sometimes the dense clusters of leaves do not appear opposite, and hide the stipule scars. A few Rubiaceae without illustrations have spines, including Chomelia spinosa and Rosenbergiodendron formosum (the latter used to be considered Randia); Guettarda foliacea (which is illustrated) occasionally has spines. Eight additional Randia species are found in Panama. Most, but not all, have small, sharp spines; all have fruits of the same size and shape. There are wet and montane forest species, and one in dry areas.

Rudgea pittieri

Rudgea pittieri

Rudgea pittieri (borojocito, fruto de mono). A

small tree of cloud-forest edge. Leaves are opposite, thick, shiny above and below, and rounded at the tip. Stipules are persistent and divide at the tip

into many tiny, pointed fingers. Flowers are white, with short tubes, held in dense, terminal heads. Fruits are large berries that turn yellow when mature. Distribution: Restricted to c. Panama, where found in lower montane and wet lowland forests of the Caribbean slope, usually along the forest edge or in clearings. Recognition: See Cosmibuena macrocarpa, which also has thick, glossy leaves. Seven more Rudgea species are found in Panama. Several are very rare, but two are fairly widespread in montane or wet forest. They do not generally have the thickened, rounded leaves of R. pittieri.

Rubiaceae

Randia armata

Rudgea pittieri

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410

Rutaceae

Warszewiczia coccinea

Warszewiczia coccinea

Warszewiczia coccinea (sanguinaria, cresta de gallo, orinera). A medium-sized tree with conspicuous red displays. Leaves are opposite, narrow, smooth and shiny. Stipules are triangular, with pointed tips, and are twisted along their length. Flowers are red, tiny, on long spikes made conspicuous by bright red, round calyx lobes that are much larger than the flowers. Distribution: In lowland wet forest in e. Panama and throughout Costa Rica. Occurs around Gamboa near the Canal, though not common. Usually seen along roads, not inside the forest. Recognition: Conspicuous when the red floral spikes are present, and they often are. Will typically be overlooked when not flowering, because the leaves are rather nondescript, though narrower than

those of most Rubiaceae. The twisted stipules are unusual, though two nonillustrated genera (Chimarrhis and Ferdinandusa) are similar. An interesting feature of the Rubiaceae is the large number of genera. Other big families like Annonaceae, Myrsinaceae, and Myrtaceae have many species in a couple huge genera. But Rubiaceae, like Fabaceae, have many genera. This should indicate to you that the species can be distinguished, but it takes a lot of work. One day there will be a guide covering the entire family, and you will not have to let any go as unidentified. Here are the Rubiaceae genera we did not cover, with species number in each: Agouticarpa (two), Allenanthus (one), Appunia (one), Bathysa (one), Bertiera (two), Borojoa (three), Chimarrhis (two), Chiococca (one), Chione (three), Chomelia (seven), Cinchona (one), Condaminea (one), Deppea (one), Elaeagia (five), Ferdinandusa (one), Gonzalagunia (four), Hillia (one), Hippotis (two), Ixora (two), Joosia (one), Machaonia (two), Morinda (three), Raritebe (one), Rogiera (one), Rondeletia (two), Rosenbergiodendron (one), Rustia (three), Simira (two), Sommera (one), Stachyarrhena (one), and Wittmackanthus (one).

Hortia colombiana

Rutaceae

Hortia colombiana

The citrus family is a moderately large group of tropical trees and shrubs, with just a few herbs, but more so in Asia and Africa. There are 1600 species worldwide, but only 350 in tropical America. The genus Citrus is from Asia, though widely planted everywhere in the world. There are 26 species native in Panama, of which we cover 5, with 4 illustrated; we also briefly describe the cultivated Citrus. The Rutaceae characters are subtle: leaves have tiny translucent spots and a citruslike odor, but the odor can be weak, and the spots are only visible with magnification when a leaf is held up to bright light. Other than those characters, they are variable. Many have alternate, compound leaves, but there are also species with simple and even opposite leaves, and we cover each pattern. The compound-leaved genus is easy to learn, but simple-leaved ones are difficult.

Citrus sinensis (orange). Not illustrated. Native to

Asia and probably an ancient hybrid. The leaves are

not at all like those of the native Rutaceae, being simple, small, shiny, and having winged twigs. When crushed, all parts have a powerful, citruslike odor. Distribution: Planted in farms, yards, and towns, and can occasionally be found inside forest where there were once farms. They do not, however, spread on their own into forest. Recognition: The odor is the key. Key lime (C. aurantifolia) and bitter orange (C. aurantium) are also planted. The simple-leaved Fabaceae, Swartzia (Papilionoideae), has leaves similar to those of Citrus, but no odor.

Hortia colombiana (limón de montaña). A tall wet-forest tree. The bark is light colored, corky, and fissured. Leaves are clustered near the ends of branches, simple, alternate, long, thick and glossy, widest close to the rounded tip. There are tiny translucent spots visible through a hand lens. Flowers are small and pink, in dense heads. Fruits are large berries that turn yellowish when mature. Distribution: Known only in Caribbean slope wet forest of Panama, not in Costa Rica. Fairly common in cloud forests at Cerro Jefe and Cerro Azul near Panama City. Recognition: The leaves are nothing like those of Zanthoxylum, resembling much more those of Pouteria (Sapotaceae), Laxoplumeria (Apocynaceae), or Anacardium (Anacardiaceae). Both Pouteria and Laxoplumeria have thick, glossy leaves, but both produce lots of white latex when broken (Hortia has no latex). Leaves of Anacardium are not thick like those of Hortia. See also Ternstroemia (Ternstroemiaceae).

Rutaceae

Warszewiczia coccinea

Hortia colombiana

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412

Rutaceae

Zanthoxylum acuminatum

Zanthoxylum acuminatum

Zanthoxylum acuminatum (arcabú, tachuelo, largarto). A medium-sized forest tree. The trunk has blunt, cylindrical woody spines fairly evenly dispersed. Leaves are very long, compound, with many leaflets (21 or so), including a single terminal leaflet. The leaf rachis has tiny, sharp spines, and the leaves have small clear dots (pellucid dots), visible when viewing through a hand lens against light. Leaflets are toothed and asymmetric, with the basal half narrower than the outer half, and smell strongly of citrus when crushed. Flowers are tiny, green, in dense, ter-

minal heads. Fruit are capsules that split open to reveal single, shiny seeds. Distribution: Mostly seen in the Canal Area, where it is very common. Otherwise sparse but widespread in lowland and lower montane forests. Not a gap-dependent species, and occurs in forest understory, growing slowly. Recognition: All Zanthoxylum have long, compound leaves, and most have spines on the leaves and trunk; all have pellucid dots in the leaves. The easiest distinction is the form of trunk spines, which vary in shape and number. Z. acuminatum has conical spines that are fairly evenly dispersed on the trunk. Its strongly fragrant leaves and tiny, sharp leaf spines are also good characters. Leaves of Zanthoxylum can be easily confused with those of Spondias mombin (Anacardiaceae), but leaves of Spondias lack spines and have a turpentine-like odor when crushed. See also Mosquitoxylum and Astronium (both also Anacardiaceae), several Trichilia (Meliaceae), plus Talisia (Sapindaceae), all with long, compound leaves.

Zanthoxylum ekmanii

Zanthoxylum ekmanii

ognition: Nearly always identified by its flat spines, which are quite unlike the conical or rounded spines of the other Zanthoxylum in the Canal Area. All the Zanthoxylum have similar leaves.

Zanthoxylum panamense (arcabú, tachuelo, largarto, palo de la cruz). Not illustrated. A tall forest

Zanthoxylum ekmanii (arcabú, tachuelo). A tall

tree of forest clearings. The trunk is covered with large woody spines that are flattened from top to bottom. Leaves are long and odd-compound, spiny, with many leaflets and pellucid dots. The leaf odor is weak. Flowers and fruits like those of Z. acuminatum. Distribution: Common at Barro Colorado I. and Soberania, otherwise just a few widely scattered records. Only found in small gaps in mature forest. It is one of the fastest growing trees in the area, reaching 30–50 cm in trunk diameter within 10–15 years. Rec-

tree. The trunk has large, cylindrical spines densely packed at the base. They often fall off, so larger trees may have few. Leaves are alternate, long, odd-compound, with many leaflets alternating (or subopposite) along the rachis. There are small, very sharp spines on the petiole and leaf rachis. There is not much odor, but the leaves have translucent dots. Flowers and fruits are like those of Z. acuminatum. Disribution: Widespread, but only common near the Canal. This species is mostly inside the forest, not edge or in gaps. Recognition: Usually known by the trunk spines being congested near the ground; in other Zanthoxylum, the spines are more or less evenly spread up and down the trunk.

Rutaceae

Zanthoxylum acuminatum

Zanthoxylum ekmanii

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Salicaceae

Zanthoxylum setulosum

Zanthoxylum setulosum

Zanthoxylum setulosum (arcabú, tachuelo). A

medium-sized forest-edge tree. Trunk spines are evenly dispersed, somewhat flattened side-to-side; they look like limpets. Leaves are alternate, odd-compound, very long and with many leaflets, the entire leaf up to 1 m in length. The leaf rachis and branches have sharp spines. Leaflets have translucent points, but are not fragrant when crushed. Flowers and fruits like those of others in the genus. Distribution: An edge species, seldom inside the forest. Mostly on the Pacific slope, including the driest zones of Panama

and Costa Rica. Very common along Pipeline Rd. Recognition: See the other Zanthoxylum species. This one has spines that are flattened (slightly) sideto-side, not top-to-bottom as in Z. ekmanii. Eight more Zanthoxylum species are found in Panama. Most are montane, though a couple are too rare to know. All have compound leaves, but some Zanthoxylum in montane areas lack spines. Zanthoxylum is so numerous that it is easy to get the impression it is the Rutaceae of Central America, but there are many more genera in the family: Amyris (three species), Conchocarpus (two), Desmotes (one), Esenbeckia (two), Galipea (two), Peltostigma (one), Raputia (one), Stauranthus (one), Ticorea (one), and Toxosiphon (one). None are common and some are very rare. Several of the other genera have compound leaves, and several simple, so you must rely on the odor and the pellucid dots to get the family correct.

Banara guianensis

Salicaceae

(Formerly Flacourtiaceae) Banara guianensis

This is the willow family, and it includes willows and poplars of the far north. But it also includes a larger group of tropical trees and shrubs. All told, there are 1200 species in the world, and about 290 in tropical America. Combining the northern and the tropical groups is a recent change, and most tropical botanists will continue to think of the tropical group as Flacourtiaceae. It has long been known, however, that the flacourts are close to willows. Panama has 39 species of Salicaceae, and we cover 12, with 11 illustrated. Salicaceae always have alternate leaves, and most have teeth. In lowland tropical rainforest, teeth are unusual, so this is a very good clue. Most tropical Salicaceae, and all those we describe here, have regularly spaced leaves in a flat plane, called

distichous. Any tree you find with toothed leaves in this distichous arrangement you should first think of as Salicaceae (and subsequently the closely related Lacistemataceae). In addition, most have tiny clear spots in the leaves (pellucid dots) that can be seen with magnification if the leaf is held up to light. The spots are very subtle and hard to see, but can be a useful clue.

Banara guianensis (corta lengua, pica lengua). A

treelet of open areas. The stem is short and leaning, and branches are long and thin. Leaves are alternate, toothed, round at the base and pointed at the tip, distichous, and have a pair of small glands just above the base. Flowers are small and white to yellowish. Fruits are berries that turn purple or black when mature. Distribution: Disturbed areas, secondary forest, especially in the rural Pacific region. Occasionally appears in old-growth forest, and remained a mystery in the old growth at Barro Colorado I. because no one recognized the few individuals that were occasionally found. Recognition: Looks just like typical Salicaceae, and closely resembles several Casearia. The leaf glands are the clue, because no Casearia has them. Note also differences in flowers and fruits.

Salicaceae

Zanthoxylum setulosum

Banara guianensis

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Salicaceae

Casearia arborea

Casearia arborea

Casearia arborea (corta lengua, pica lengua,

manga larga). A medium-sized forest tree. The trunk is straight and branches high. The bark is light yellowish. Branches are horizontal, thin and long.

Leaves are alternate, toothed, round at the base and pointed at the tip, regularly spaced and arranged in a flat plane, with pellucid dots. Flowers are white to greenish, in small groups on branches between the leaves. Fruits are three-valved capsules that split open to reveal reddish orange seeds. Distribution: In moist to wet and montane forests throughout. It is a species of natural clearings, so juveniles are usually found only in openings; adults are quite common. Recognition: The leaves fit precisely the Salicaceae mold. Even in big trees, the very long branches with leaves regularly spaced along the whole length are easy to see. Laetia procera has a very similar look. The other common Casearia are not as tall.

Casearia arguta

Casearia arguta

Casearia arguta. A small tree of forest edges. Leaves are alternate, toothed, distichous, much longer than they are wide, and straight-sided. Flowers are white, in clusters along the branches. Fruits are capsules, also along branches. Distribution: Mostly a Pacific slope species, and occurs in the driest zones, mainly at forest edge and roadsides. Recognition: The long, narrow leaves are like those of Laetia procera, but the latter is a tall tree.

Salicaceae

Casearia arborea

Casearia arguta

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Salicaceae

Casearia commersoniana

Casearia commersoniana

Casearia commersoniana (corta lengua, pica

alternate, in a flat plane, with pellucid dots. Teeth are conspicuous but sparse and dull. Flowers are white to green, along branches. Fruits are small capsules that darken to purple when mature. Distribution: All types of forest, from dry to wet lowlands and sometimes lower montane. Very common and widespread around the Canal. Recognition: Other Casearia have teeth densely packed along the leaf margin, but in C. commersoniana, teeth are widely spaced. Branches are also somewhat shorter, with fewer leaves.

lengua, mauro, yerce). A small forest tree. Leaves are Casearia sylvestris

Casearia sylvestris

Casearia sylvestris (corta lengua, pica lengua). A

medium-sized tree of regrowth. The trunk typically forks at a low height. Branches are long and thin. Leaves are simple, alternate, distichous, with pellucid dots, teeth either small and inconspicuous or lacking.

Flowers are white to greenish, in clusters along branches. Fruit capsules open to reveal seeds with red arils. Distribution: Very widespread, in all types of forest, generally in secondary forest and regrowth, though also in old growth. Seen often throughout the Canal Area. Recognition: The long branches with regularly spaced leaves resemble those of the rest of the family. This species has faint or even no teeth on leaves, and thus might look like Annonaceae, but the pellucid dots will distinguish it. Branches might be mistaken for compound leaves because they are long and horizontal and leaves are small. Casearia has nine more species in Panama. Several are from dry regions, others are montane, and a few are very rare. The extent of leaf teeth is variable, and those with few teeth are more difficult to recognize.

Salicaceae

Casearia commersoniana

Casearia sylvestris

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Salicaceae

Hasseltia floribunda

Hasseltia floribunda

Hasseltia floribunda (parimontón, corta lengua,

raspa lengua). A small or medium-sized tree of forest edge. The trunk is frequently branched near the ground. Leaves are alternate and toothed like those of other Salicaceae, but angled relative to the petioles and thus not in a flat plane; as a caution, though, leaves are variable and sometimes even lack teeth. Three large veins emerge together from the leaf base.

At the base of each leaf there is a pair of flattened, oval glands, adjacent to the petiole where it meets the leaf. Petioles are slightly swollen at both ends. Flowers are small, white, in loose clusters at the end of branches. Fruits are small berries that turn purple when mature. Distribution: Widespread in all forest types, including lower montane. Very common around the Canal, in old-growth and secondary forest and often along wooded roads. Common along Pipeline Rd. and easy to see in flower there. Recognition: Hasseltia is not confused with its relatives in the Salicaceae, because its leaves have three major veins and are not in a flat plane. It is confused with the remarkably similar Alchornea costaricensis (Euphorbiaceae), which even has glands in the same position. In Hasseltia, the three basal veins extend toward the leaf tip; in Alchornea, the two side veins are smaller and do not extend as far toward the tip.

Laetia procera

Laetia procera

Laetia procera (manga larga). A large tree of forest clearings and edge. The trunk is straight and cylindrical, and bark is dark brown with black spots. Branches are horizontal, long, and thin, with many leaves per branch. Leaves are simple, alternate, regularly spaced, in a flat plane, densely toothed along

the margins, long and straight-sided. Flowers are small, white, in clusters along branches. Fruits are capsules that turn red and split open. Distribution: Mostly on the Caribbean half of the isthmus in Panama and sparse in Costa Rica, never particularly common. Typical of clearings and roadsides, and some tall trees have branches arching over Pipeline Rd. In the forest, juveniles invade gaps and are rare in the shade. Recognition: The very long branches, packed with regularly spaced leaves, and the long, straight-sided leaves, are easy to spot along roads or around forest gaps. Leaves are like those of Casearia arguta, but that species is a treelet. Casearia arborea is a tall tree, but has different leaves. L. thamnia has much smaller leaves.

Salicaceae

Hasseltia floribunda

Laetia procera

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Salicaceae

Laetia thamnia

Laetia thamnia (conejo, conejo colorado, palo blanco). A medium-sized forest tree. The trunk is

sometimes teeth are absent. Trees of wetter forest have fewer teeth. Flowers are white with yellow stamens, and are quite a bit larger than others of the family. They are held singly along branches. Fruits are also large and open to reveal seeds covered in yellow-orange arils. Distribution: In lowland wet to lower montane forests throughout, but not particularly common. Recognition: When toothed, it should make you think Salicaceae, and then the shiny green leaves are much different from those of Casearia and other Laetia. Without teeth, it looks like Oxandra longipetala (Annonaceae).

straight, cylindrical, and bark is white or yellowish. Branches are long. Leaves are alternate and regularly spaced, dark shiny green, usually blunt-toothed, but

One more Laetia species is known, but it is rarely seen and only in far e. Panama.

Laetia thamnia

Ryania speciosa

Ryania speciosa

Ryania speciosa (corta lengua). An understory tree. Leaves are alternate, regularly spaced and in a flat plane, but lack teeth. Secondary veins are straight

and parallel, impressed above. Most leaves have large swollen and cup-shaped insect galls. Flowers are large, with white petals in a star form and pink anthers. Fruits are large, spherical capsules that turn red when mature. Distribution: In wet forests of the Caribbean slope and lower montane sites in Panama, and lowland wet forest in Costa Rica. Common near the Canal. Always inside mature forest. Recognition: The galls are the best character; near the Canal, leaves are always infected and no other species has anything like them. Because it lacks teeth, the leaves look like those of Annonaceae, and Annona might come to mind.

Salicaceae

Laetia thamnia

Ryania speciosa

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Salicaceae

Tetrathylacium johansenii

Tetrathylacium johansenii

Tetrathylacium johansenii (palo de chancho,

broadest near the tip, with an unequal base. Flowers are small and white, held on spikes, and many spikes are clustered together at branch ends. Distribution: Widely seen throughout the Canal Area; otherwise, seldom seen. In mature forests, and often in swampy areas or along rivers. Recognition: The leaves match the Salicaceae pattern, but are broadest above the middle, and this is a good character. Look for the sprouts too, which seem to appear on most big trees and provide leaves within easy reach even if the tree is tall.

tresses extending into long superficial roots. From all around the base of the trunk, even the roots, small sprouts grow upward, sometimes dozens on one tree. Leaves are alternate, toothed, distichous, usually

A second species of Tetrathylacium is known from wet forests of c. Panama and sw. Costa Rica. It has similar but larger leaves, and does not form trunk sprouts.

pantano). A tall forest tree. The trunk has small but-

Xylosma chlorantha

Xylosma chlorantha

Xylosma chlorantha (cachos de venado). Not il-

lustrated. A spiny treelet. The trunk is covered with many-branched spines. Leaves are simple, alternate, with obvious teeth; there are no pellucid dots. Flow-

ers are yellow and held in groups along the branches. Fruits are fleshy, oval, turn yellow or red when mature. Distribution: Sporadically distributed along the Pacific coast and sometimes in wet and lower montane forest, but never common. In the Canal Area, it occurs in lowlands near Panama City, the Caribbean, and in mountains, but is not numerous. Recognition: The branched spines are very distinctive. A species of Casearia, C. aculeata (not illustrated), has spines on the trunk, but they are not forked. Eight more species of Xylosma are found in Panama, all treelets with branched trunk spines and toothed leaves.

Zuelania guidonia

Zuelania guidonia

Zuelania guidonia (cagajón, árbol caspa, dan-

druff tree, caraño). A medium-sized tree of dry areas. The trunk is straight and cylindrical, and bark is brown with many white lenticels. When rubbed, the lenticels flake off as little white scales, like dandruff. Leaves are alternate, regularly spaced, in a flat plane, finely toothed or not at all, with the base asymmetric; secondary veins are numerous and parallel. The underside is pubescent, with reddish veins. During the dry season, leaves turn yellow before dropping, and most trees have a few yellow leaves mixed in with

green. Flowers are white, in clusters along branches. The fruit is a spherical capsule that splits open to reveal seeds with orange arils. Distribution: Mostly on the Pacific slope, where it occurs in the driest zones of c. Panama and nw. Costa Rica. We see it occasionally along the Caribbean slope in Panama, too. Typically in secondary forest, edge, and pasture. Common near the Canal. Recognition: Teeth are very fine and might be missed, then suggesting Annona spraguei (Annonaceae) or Castilla elastica (Moraceae), which grow in the same habitat and have similar leaves. Compared to other Salicaceae, the reddish pubescence and asymmetric leaf base are distinctive. There are six genera in the Salicaceae in Panama that we have not treated: Homalium, Lunania, Macrohasseltia, Pleuranthodendron, Prockia, and Salix, all with a single species except Lunania, which has two. Except Lunania, all have toothed leaves and should be recognized as Salicaceae.

Salicaceae

Tetrathylacium johansenii

Zuelania guidonia

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Sapindaceae Allophylus psilospermus

Sapindaceae Allophylus psilospermus

One of the major tropical forest families, with 2200 species worldwide and 950 in tropical America. The largest number, however, are lianas, with trees accounting for only about a third of the species. Some Sapindaceae are found in subtropical and temperate areas, in fact, in the new angiosperm classification, the maples (genus Acer) are part of Sapindaceae, meaning that the family is now a major temperate zone group as well. The well-known Asian fruits litchi (genus Litchi) and rambutan (Nephilium) are Sapindaceae, as is the Brazilian caffeine drink guarana (Paullinia), but no Panamanian species is used for fruit. The Panama tree checklist includes 26 species of Sapindaceae, and we cover 11, with 9 illustrated. Sapindaceae is one of the major compoundleaved families, along with Anacardiaceae, Burseraceae, Fabaceae, and Meliaceae, and they are usually very easy to recognize. The base of the petiole is brown and swollen where it meets the branch, flattened above or on the sides; this trait is shared with Anacardiaceae, Burseraceae, and Meliaceae, but Fabaceae have a cylindrical, green petiole base. Leaflets of Sapindaceae are alternate

along the rachis, and the rachis extends a short distance beyond the last leaflet to an aborted tip. The latter is an excellent character, unlike any other family but found in most (not quite all) trees of the Sapindaceae. In the most common tree genera, the alternate leaflets are toothed, adding another easy and distinctive feature. Flowers are tiny and white, in dense terminal clusters, and not distinctive. Many of the species have capsular fruits that split open to reveal seeds with small arils attractive to animals.

Allophylus psilospermus (esquitilla, esquitillo).

A trifoliate understory forest tree. The bark is creamcolored. Leaves are alternate, compound, always with three leaflets that are slightly toothed; they are smooth and hairless. The petiole is slightly swollen and brown where it leaves the branch. Flowers are tiny, white, packed on spikes that themselves are in a dense cluster. Fruits are fleshy, not capsules. Distribution: Widespread, including lower montane sites and the dry Pacific slope plus moist to wet forest, but sporadic and not common. Only in forest. Recognition: Trifoliate compound leaves with teeth are rare. See Crateva (Capparaceae), but its leaflets are not toothed. Hevea brasiliensis (the rubber tree [Euphorbiaceae]) has trifoliate leaves on very long petioles, but is not native and is only seen in towns. Two other Allophylus species are found in Panama, both with trifoliate leaves; in one, they are barely toothed. The other, A. racemosa, is in very dry areas and has fuzzy, clearly toothed leaflets.

Cupania cinerea

Cupania cinerea

Cupania cinerea (gorgojero, gorgojo blanco). A medium-sized tree of forest edge. Larger trees have small buttresses at the base, and bark and branchlets have white lenticels. Leaves are alternate, compound, usually with seven alternating leaflets. The terminal

leaflet is angled to one side, and at its base is a small aborted rachis tip. Leaf undersides are whitish due to a dense coat of hairs. Flowers are tiny, white, in pyramid-shaped, many-branched terminal clusters. Fruits are three-parted capsules that open to reveal three seeds. Distribution: Widespread around the Canal, and common there along roads on the Pacific half of the isthmus. Seldom seen elsewhere. Recognition: The genus Cupania is very easy to learn, with toothed alternate leaflets and the little “coupon” at the end of the rachis. Moreover, the various species are usually easy to tell apart: C. cinerea is the one with whitish leaf undersides. A caution though, as juveniles, most Cupania can have simple leaves.

Sapindaceae

Allophylus psilospermus

Cupania cinerea

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Sapindaceae

Cupania latifolia

Cupania latifolia

Cupania latifolia (gorgojero, gorgojo). A medi-

um-sized forest-edge tree. Trunk is straight but fluted, slightly swollen into irregular buttresses at the base. The bark has lenticels. Leaves are alternate, compound, with alternating leaflets (though basal leaflets can be opposite), rounded at the tip; veins are

impressed above, so leaflets are bullate. There is a terminal leaflet, but it is angled to one side; at its base is a short stalk angling away from the leaflet, the Cupania “coupon.” Leaflets have somewhat toothed margins, but teeth are wavy and irregular. The base of the petiole is swollen, brown in color, and flattened. Flowers are tiny and white. Fruits are threeparted capsules, and the black seeds have a yelloworange aril. Distribution: Almost exclusively in the Canal Area, where it is widespread at both lowland and lower montane sites. Not common inside the forest, but can be seen along roads in wooded areas. Recognition: Easy to know as Cupania. This species has dark leaflets, rounded at the tip, with blunt and irregular teeth.

Cupania rufescens

Cupania rufescens

Cupania rufescens (candelillo, gorgojero, gorgojo). A medium-sized forest-edge tree. Leaves are alternate, compound, and the leaflets are alternate along

the rachis. The final leaflet is angled slightly to one side, appearing terminal; it has a short stalk adjacent to it. All leaf parts and small branchlets are covered with dense, reddish hairs. Flowers and fruits are like those in C. latifolia. Distribution: Very common along roads in the Canal Area from Panama City to Pipeline Rd.; elsewhere widespread in lowlands. Not common inside the forest. Recognition: Often seen and easy to recognize. None of the other Cupania or any other Sapindaceae have anything like the red hairs. Might be confused with Inga goldmanii (Fabaceae—Mimosoideae), which has red hairs but opposite leaflets.

Sapindaceae

Cupania latifolia

Cupania rufescens

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Sapindaceae

Cupania scrobiculata

Cupania scrobiculata

Cupania scrobiculata. Not illustrated. A medium-sized forest tree. As a sapling or small tree, the trunk is ribbed or fluted. Otherwise, it is a typical Cu-

pania with compound, alternate leaves and alternating, toothed leaflets, plus the aborted rachis tip. Veins are impressed above, leaflets pointed. Distribution: We see it all over the Canal Area, though oddly it skips over Barro Colorado I. and has never been seen there. Elsewhere barely recorded. Common inside forest, not forest edge. Recognition: In the understory, note the fluted stem of saplings, like that of Tachigali (Fabaceae—Caesalpinioideae). Leaves are shaped like those of Cupania seemannii but have teeth. In color and texture they are more like those of C. latifolia, but with impressed veins and pointed (not blunt) leaflets.

Cupania seemannii

Cupania seemannii

Cupania seemannii (gorgojero, gorgojo). A small

or medium-sized forest tree. Trunk is irregular and often has many sprouts at its base. Leaves are alternate, compound, with alternating leaflets. The terminal leaflet is angled to one side, and at its base there is a

small point angling in the other direction. Leaflets are untoothed. The petiole has a swollen base, brown and flattened. Distribution: Yet another species easy to find in the Canal Area but seldom seen elsewhere. Fairly common inside mature forest. Recognition: Unlike the other Cupania, leaflets lack teeth, and thus resemble those of some Matayba. But it does have the little “coupon” at the end of the leaf. Four more Cupania species are known in Panama, all with the usual Cupania traits. So far we have covered several largely localized to the Canal Area; C. guatemalensis is a dry-zone species well known in nw. Costa Rica plus c. Pacific Panama. Two of the species are very localized and poorly known.

Matayba apetala

Matayba apetala

Matayba apetala (gorgojero, gorgojo). A medi-

um-sized wet-forest tree. The trunk is slightly fluted, and bark black. Leaves are usually opposite, though also alternate on some occasions. They are compound, with many narrow, shiny leaflets lacking teeth, and the leaflets are alternate to subopposite along the rachis. (So leaves are mostly opposite, leaflets mostly alternate, but both traits are variable.) The

end leaflet is turned well to the side, and there is a little pointed swelling at its base. Distribution: Widespread in the Caribbean half of the Canal Area; otherwise barely known. We recently found in Bocas del Toro, suggesting it is a widespread Caribbean lowland species. Recognition: Very unusual among Sapindaceae, this species has opposite leaves, but it should still be recognized to family. Matayba is closely related and similar to Cupania; both groups have the end leaflet angled to one side and a point at the end of the rachis (shorter in Matayba). Both genera also have alternate leaflets, though in Matayba they tend toward subopposite. Also, nearly all Cupania have toothed leaflets, whereas many Matayba do not. Once recognized as Sapindaceae, M. apetala is known from its narrow, shiny, numerous, untoothed leaflets.

Sapindaceae

Cupania seemannii

Matayba apetala

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Sapindaceae

Matayba scrobiculata

Matayba scrobiculata

Matayba scrobiculata (laso, laso prieto, matillo). A medium-sized tree of the dry zone. The trunk is slightly fluted, and the bark has conspicuous orange patches. Leaves are alternate, compound; leaflets are subopposite or slightly alternate, blunt at the end, and have toothed or wavy margins. One of the final pair of leaflets points forward, appearing like a terminal leaflet, and there is often a small projection at the

base of this forward-pointing leaflet. On the underside, there are small pits in the axils of the main vein. Flowers and fruits are like those of Cupania. Distribution: Found only on the Pacific half of the isthmus. Common in and around Panama City and in farmland and shrubland of the driest section of Panama. One record in Costa Rica. Recognition: Readily known as Sapindaceae, this species could be mistaken for Cupania latifolia, which has similar wavy-margined, blunt-tipped leaflets. For further confusion, note that Cupania scrobiculata resembles M. scrobiculata, and both are found on the Pacific slope. M. apetala is restricted to wetter areas and has much narrower leaves than M. scrobiculata. Three other Matayba species are found in Panama. M. glaberrima is another dry-zone species of Panama, rare in Costa Rica. The other two are very rare.

Sapindus saponaria

Sapindus saponaria

Sapindus saponaria (jaboncillo, soapberry). A medium-sized tree of open areas. The bark is yellowish or cream-colored. Leaves are alternate, compound, with leaflets opposite (or nearly so) along the rachis; there is no terminal leaflet, but one of the last pair is angled forward. Leaflets are asymmetric, the outer half wider than the inner half, and tend to curve backwards. The leaf rachis is ridged or even slightly winged between leaflets. The base of the petiole is swollen and flattened. Flowers are small, white, in dense clusters at the end of branches, conspicuous from a distance where it occurs along

roads. Fruits are medium-sized, spherical berries, in groups of two or three. When mature, they are yellow and covered in a mucilaginous membrane. Distribution: A species of the dry Pacific slope in c. Panama and nw. Costa Rica, usually not inside the forest but common in open areas or along forest edge. Recognition: Like other Sapindaceae in the arrangement of the last pair of leaflets, but the deflection of the end leaflet is weak. Sapindus has more leaflets than other Sapindaceae, and the leaflets are mostly opposite. All together, Sapindus may not immediately jump out at you as Sapindaceae. The winged rachis is reminiscent of Inga (Fabaceae—Mimosoideae), but all Inga have rachis glands, and Dipteryx (Fabaceae—Papilionoideae), which also has asymmetric leaflets. Recall also that Fabaceae have a cylindrical swelling at the base of the petiole, unlike the flattened swelling in Sapindaceae. Simarouba amara (Simaroubaceae) has leaflets similar in size and shape to those of Sapindus, but they are bright shiny green. Fruits of Sapindus are used for making soap, as the name suggests.

Sapindaceae

Matayba scrobiculata

Sapindus saponaria

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Sapotaceae

Talisia croatii

Talisia croatii

Talisia croatii. Not illustrated. An understory treelet with an unbranched stem and long leaves with many leaflets radiating off it horizontally, generally bunched near the top. Leaflets are smaller and more numerous than in T. nervosa. The top of the stem in

T. croatii has a very unusual, leafly stipule, like a tiny, compound leaf. Flowers are tiny, white, and in spikes that are clustered at the top of the stem. Distribution: Mostly known in the Canal Area, but there are a few other records, including sw. Costa Rica. Recognition: Unlike other Sapindaceae. The even-numbered, opposite leaflets resemble those of many Fabaceae, and it is easily confused with Tachigali versicolor (Fabaceae—Caesalpinioideae), which has similar leaf arrangement and form, plus the same sort of weird, divided stipule. Recall that Fabaceae have green, cylindrical petiole bases, whereas Sapindaceae have brown, flattened petiole bases. Compared to T. nervosa, this species has smaller and more numerous leaflets.

Talisia nervosa

Talisia nervosa

Talisia nervosa (mamón de montaña). A treelet of forest understory. The stem is straight and unbranched, with long leaves radiating off it horizontally and generally bunched near the top. Leaves themselves are alternate, very long but compound, with opposite leaflets. The leaflets are long, and they get larger and longer toward the tip. This species lacks the compound stipule of T. croatii. Flowers as in T. croatii. Distribution: In c. Panama on the Caribbean

slope, plus wet forests of sw. Costa Rica. Recognition: See T. croatii and also Tachigali versicolor (Fabaceae—Caesalpinioideae), whose saplings resemble those of Talisia. Panama has six more Talisia species, but five are barely known. One of them, T. princeps, has been confused with T. croatii, but both species are identified in Panama. Sapindaceae has five more genera in Panama: Billia, Blighia, Dilodendron, Dodonaea, and Pseudima. All have one species except Billia, which has two. Dodonaea is completely different from others in the family in having simple leaves. Billia has opposite, trifoliate leaves, and used to be in the horsechestnut family, Hippocastanaceae. The other three have compound leaves, and Pseudima looks very much like Matayba and Cupania.

Chrysophyllum argenteum

Sapotaceae

Chrysophyllum argenteum

One of the major families of tropical forests, nearly all of which are trees or treelets and almost all are tropical. There are 1100 species in the world and about 400 in the American tropics. Several American Sapotaceae produce edible fruits, and many of these are either collected from natural forest or grown in plantations. In Panama, the fruits of Chrysophyllum cainito and Manilkara zapota are readily available. It is the family of chicle (Manilkara), originally the basis of chewing gum. All told, 57 tree species in the Sapotaceae are native in Panama, including 31 in the

genus Pouteria. We cover 13 Sapotaceae here, with 12 illustrated. All Sapotaceae produce white latex that drips from broken parts, as in Apocynaceae, many Moraceae, and some Euphorbiaceae. In many Sapotaceae, leaves are clumped at the ends of branches and arranged in spirals, but some have regularly spaced leaves in a flat plane. In addition, many Sapotaceae have smooth leaves with closely spaced venation, but again, this is not universal in the family. For any tree with a lot of latex, Sapotaceae should be among your first thoughts; if leaves are clustered and spiraled, this family becomes a prime candidate. Flowers of Sapotaceae are tiny, borne along branches. Fruits are big, round, and have sweet, fleshy pulp and large, shiny seeds bearing a longitudinal scar.

Chrysophyllum argenteum (caimito de mono).

A tall forest tree. The trunk can have small buttresses at the base. Twigs and petioles have light, silvery pu-

Sapotaceae

435

Talisia nervosa

Chrysophyllum argenteum

bescence, and petioles are grooved above. Leaves are simple, alternate, regularly spaced and in a flat plane, dark shiny green above and light shiny green below. Any broken part produces much white latex. As in many species with latex, the leaves have a slightly thickened, stiff feel. Flowers are small, yellowish, in short-stalked clusters along the branches. Fruits are fleshy and soft, plum-sized, turning purple when mature. Distribution: In lowland moist to wet

forests throughout, never especially common. Recognition: Leaves are larger and lack the red color of the related C. cainito. Might be confused with Aspidosperma spruceanum (Apocynaceae), which also has shiny, thickened leaves with latex, but the latter has veins more closely spaced. In Pouteria, leaves are bunched at the ends of branches. Moraceae and Euphorbiaceae that produce as much latex are quite different.

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Sapotaceae

Chrysophyllum cainito

Chrysophyllum cainito

Chrysophyllum cainito (caimito). A large forest tree. Big individuals have wide crowns emerging above the surrounding canopy. The trunk is slightly buttressed or irregular when large. Petioles are grooved above. Leaves are alternate, arranged regularly along branches, in a flat plane. The undersides of the leaves are a striking, velvety, somewhat iridescent red, created by dense, soft hairs. The twigs zig-

zag between leaves and also have soft, red pubescence. Like all Sapotaceae, broken leaves, or cut bark, produce dripping white latex. Flowers are similar to those of C. argenteum. Fruits are plum-sized or larger, bluish, with a soft flesh that has a spicy flavor. Distribution: Found in the dry zone of Panama, but also widespread in lowland moist to wet forests. Frequent in forests near Panama City and on Barro Colorado I., where it is conspicuous but not especially numerous in the old growth. Often found outside the forest and in agricultural areas. Recognition: With a leaf in hand, this species cannot be confused. Luehea seemannii (Malvaceae—Grewioideae) is another tall tree with reddish leaf undersides, and it might be mistaken for caimito at a distance, but leaves of Luehea have much different venation and texture. See also Vismia (Hypericaceae). Caimito fruits can be purchased in large markets.

Chrysophyllum colombianum

Chrysophyllum colombianum

Chrysophyllum colombianum (mameicillo, nis-

perillo). A tall wet-forest tree. The trunk can be slightly buttressed at the base. Twigs are slightly hairy. Leaves are long, narrow, shiny green, bunched toward branch ends. Veins are yellowish and raised be-

low. The petiole is swollen at the base where it meets the leaf. Broken leaves exude ample white latex. Flowers are like others of the genus; fruits are fleshy and orange. Distribution: Restricted to wet and lower montane forests on the Caribbean side of the isthmus. Recognition: The leaves are bunched and the petiole swollen, making this easily confused with several Pouteria rather than other Chrysophyllum. Five more Chrysophyllum species are found in Panama, all large trees with latex and shiny leaves. Some have regularly spaced leaves, but others have leaves bunched at the branch end as in Pouteria. Two are montane, one from wet forest, and two very poorly known.

Sapotaceae

Chrysophyllum cainito

Chrysophyllum colombianum

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Sapotaceae

Ecclinusa lanceolata

Ecclinusa lanceolata

Ecclinusa lanceolata (mameicillo, nisperillo, cai-

mito de montaña). A tall wet-forest tree. The trunk can have small buttresses at the base. Leaves are simple, alternate, shiny, somewhat thickened, and clustered toward the ends of branches. Secondary veins

are raised on the leaf underside, numerous and parallel. The base of the petiole is swollen and cylindrical. Small brown stipules sometimes remain on branches where the petiole meets. All parts drip white latex when broken. Flowers and fruits are similar to those of others in the family. Distribution: Cloud forest and wet forest in c. Panama. Quite common at Cerro Jefe. Recognition: Much like Pouteria in leaf clustering and the swollen petiole, and will undoubtedly often pass for a Pouteria. But the stipules are a distinction: Pouteria never has them. Be careful, though, because the stipules are often lost in Ecclinusa. A second Ecclinusa species is known only in Bocas del Toro and is quite similar in overall appearance.

Manilkara bidentata

Manilkara bidentata

Manilkara bidentata (níspero, níspero de montaña, balata). A wet-forest giant. The crown of large trees emerges above the rest of the canopy. Large trunks have only small buttresses. The bark has vertical fissures. Leaves are simple, alternate, tightly bunched at the ends of branches, shiny green above and light brownish below. Secondary veins are close-

ly spaced and parallel, but barely visible. The petiole is not swollen. All parts drip white latex when broken. Flowers are small, white to yellowish. Fruits are spherical, with a pointed tip, fleshy, and turn orange when mature. Distribution: Caribbean wet and lower montane forests of Panama but not known in Costa Rica. Sometimes big trees dominate the canopy, such as near the research crane at the Chagres River near the Caribbean Canal entrance. Recognition: The smooth leaves with latex and obscure secondary veins are clear characters, and this species is especially distinctive from the brownish tint to the leaf underside. But beware the genus Micropholis, which we do not illustrate but describe below. See also Chrysophyllum cainito, which has smaller leaves with a deeper red underside.

Sapotaceae

Ecclinusa lanceolata

Manilkara bidentata

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Sapotaceae

Manilkara zapota

Manilkara zapota

Manilkara zapota (níspero). A large dry-forest tree. Very similar to M. bidentata, but leaves are light

shiny green below, without any brownish. Distribution: In dry forests along the Pacific coast, and appears (possibly planted) on farms in rural areas. Occasionally found near the Caribbean too. Recognition: See the other Manilkara and also Micropholis, all with faint secondary venation. This is the only species of the group seen in dry forest. Two other Manilkara species are found in Panama. One is very rare. The other, M. chicle, is rare in Panama but frequent in nw. Costa Rica. It is another large tree similar to M. bidentata.

Micropholis melinoniana

Micropholis melinoniana

Micropholis melinoniana. Not illustrated. A tall, distinctive wet-forest tree. Leaves are simple, alternate, very shiny green, and with very closely spaced secondary veins that are just barely visible. Broken parts drip white latex. Distribution: Carib-

bean wet forest and lower mountains of c. Panama to Costa Rica. Recognition: The venation is just like that of the better-known and more widely distributed Calophyllum longifolium (Clusiaceae), but Calophyllum has opposite leaves. Manilkara zapota is also similar, but Micropholis has much more closely spaced secondary veins. Four more Micropholis species are found in Panama, all from wet to montane forest and generally resembling Manilkara in having obscure secondary venation. One, Micropholis venulosa, has reddish leaf undersides, similar to those of Manilkara bidentata. You might learn them as the alternate-leaved, wet-forest Calophyllum (Clusiaceae).

Pouteria fossicola

Pouteria fossicola

Genus Pouteria. One of the very difficult tree

genera in tropical America. As a group, they are reasonably recognizable, based on copious latex and cylindrical petioles that are swollen at the base and resemble little bottles. Their leaves are usually bunched toward the end of branches, and are often widest beyond the middle. Within the genus, they are diverse and difficult to identify. Thirty-one species are known in Panama, and in wet forests of the Caribbean slope, we are often

stumped and many remain unidentified. We cover five species here.

Pouteria fossicola (mamey de montaña, mamey). A tall, rare forest tree. The trunk can have small buttresses. Bark is brown and slightly corky. Leaves are simple, alternate, widest above the center, bunched toward the ends of branches, light colored on the underside. Veins are raised below and prominent. The petiole is cylindrical and slightly swollen at the base. Any broken part drips white latex. Flowers are white, in short clusters held on branches below leaves. Fruits are large, oval. Distribution: Widely known in wet and lower montane forest, but not common. Recognition: Rare so difficult to learn. Much like Chrysophyllum colombianum and Pouteria glomerata. At Barro Colorado I., sometimes confused with Alseis blackiana (Rubiaceae), which has similar venation and leaves that are widest above the middle. But Alseis has opposite leaves and lacks latex.

Sapotaceae

Manilkara zapota

Pouteria fossicola

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Sapotaceae

Pouteria glomerata

Pouteria glomerata

Pouteria glomerata (nisperillo, mameicillo). A medium-sized forest tree. The trunk is often slightly

fluted. Leaves are simple, alternate, clustered at branch ends, widest above the middle. The leaf underside is silvery. Petioles are cylindrical and swollen at the base. Any broken part drips white latex. Small flowers are in clusters on branches below leaves. Fruits are spherical, but have a short, sharp point at one end; they turn yellow when mature. Distribution: Known sparsely in wet forest, also on limestone. Recognition: The silvery underside is a good character, and distinctive near the Canal, where no other Sapotaceae is similar.

Pouteria reticulata

Pouteria reticulata

Pouteria reticulata (faldita de puta). A tall forest tree. Big trees have modest buttresses. The bark is brown and slightly fissured; small pieces peel off. Leaves are simple, alternate, narrow, shiny green,

typically with wavy margins. Leaves frequently have tiny, round, black spots across the upper surface, probably a fungus. The petiole is grooved above, not swollen as in others of the genus. Any broken part drips white latex. Flowers are tiny, greenish yellow, on branches below leaves. Fruits are oval, fleshy, and turn dark when mature. Distribution: In Costa Rica and Panama, from Pacific lowlands to lower mountains; also found throughout the Canal Area, including Barro Colorado I., where it is the only abundant Pouteria. Recognition: Where common near the Canal, the narrow, wavy leaves are easy to spot, and the latex cinches it.

Sapotaceae

Pouteria glomerata

Pouteria reticulata

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Sapotaceae

Pouteria stipitata

Pouteria stipitata

Pouteria stipitata (faldita de puta). A mediumsized forest tree. The bark is brown, slightly fissured, and peels in small pieces. Big trees have small but-

tresses. Leaves are simple, alternate, clustered at the ends of branches. There are few, widely spaced secondary veins. The petiole is densely pubescent, cylindrical, and swollen at the base. Any broken part drips white latex. Flowers and fruits are like those of P. reticulata, but yellow at maturity instead of black. Distribution: Along the Pacific slope from c. Panama through nw. Costa Rica. Known in dry forest and wet forest, but sparse everywhere. Uncommon at Barro Colorado I. Recognition: At a glance, much like the more common P. reticulata, but the widely spaced veins and pubescent petiole are distinctive.

Pouteria torta

Pouteria torta

Pouteria torta. A tall forest tree. Leaves are simple, alternate, bunched at branch ends, and somewhat rounded. Veins are yellowish and raised below. Petioles are cylindrical and swollen. Any broken part drips white latex. Flowers are like those of other Pou-

teria. Fruits are bizarre, covered with long, soft filaments. Distribution: The Caribbean slope and the Osa Peninsula. Quite common in wetter lowlands and low mountains near the Canal, including the Pipeline Rd. area. Recognition: Not very distinctive unless bearing fruit. Leaves are larger and rounder than in other Pouteria. We have only covered the Pouteria species likely to be seen around Barro Colorado I., Soberania, and Panama City. Beware that in wet forests, there are many more species, including some similar to those just described. Others resemble Chrysophyllum or even Manilkara.

Sapotaceae

Pouteria stipitata

Pouteria torta

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Scrophulariaceae Sideroxylon celastrinum

Sideroxylon celastrinum

Sideroxylon celastrinum (espino rico). A small tree of dry, open areas. The trunk usually branches near the ground, forming a wide, low crown. Branches have small but sharp spines. Leaves are small, rounded and widest above the middle. They are technically simple and alternate, but are clustered in groups (based in fascicles) and so do not really look alternate and might be mistaken for compound. Any

broken part drips white latex. Flowers are small, greenish, in short clusters along branches below leaves. Fruits are small berries that mature purple. Distribution: Known only in dry savanna country of c. Panama and nw. Costa Rica. Recognition: Completely unlike the rest of the Sapotaceae. Other spiny savanna trees lack latex. Four more Sideroxylon species are found in Panama, two in drier areas and two rather uncommon in mountains of the east or west. The others do not have such small leaves, nor do they have fascicles. S. persimilis in dry areas also has sharp spines. Three more genera of Sapotaceae are found in Panama, each with one species: Chromolucuma, Elaeoluma, and Sarcaulus. They have latex and bunched leaves, and are easily confused with Pouteria.

Scrophulariaceae (Formerly Loganiaceae and Buddlejaceae) Buddleja americana

Buddleja americana

A very large family, most of which are temperate herbs, diverse and well-known in North America and Europe. There are 4000 species worldwide and 750 in the American tropics, but those are mostly herbaceous, so as a tropical tree family Scrophulariaceae is insignificant. We cover the one treelet genus known from Panama and Costa Rica. It has a confusing taxonomic history, recently being placed in the Loganiaceae and then its own family, the Buddlejaceae, before landing in the much larger Scrophulariaceae. Scrophulariaceae generally have opposite, toothed leaves and flowers with extended lower lips, like those of Lamiaceae and Bignoniaceae. Buddleja

might also be confused with the Rubiaceae because of the leafy stipule between each leaf pair that falls to leave a scar, but Rubiaceae do not have teeth.

Buddleja americana. A treelet of forest edge in the mountains. Leaves are opposite, heart-shaped, and distinctly brown underneath from fine hairs. Stipules cross between leaves and usually fall off, just like in Rubiaceae. Flowers are tiny, in dense heads. Distribution: In mountains from c. Panama through Costa Rica, where it can be common along roads and in other open areas. Recognition: At a glance, this looks like Vismia baccifera (Hypericaceae), but Vismia has latex. The twig scar between leaves resembles the key trait of Rubiaceae; Arachnothryx buddleioides is the only Rubiaceae with brown leaf undersides. Three other species of Buddleja are known in Panama. Two are found in mountains from w. Panama through Costa Rica, and both have distinctive brown or white leaf undersides. The third species has just one out-of-place specimen in w. Panama, and is otherwise known only in South America.

Scrophulariaceae

Sideroxylon celastrinum

Buddleja americana

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Simaroubaceae

Simaroubaceae Quassia amara

Quassia amara

A small, mostly tropical family of trees and shrubs, with 130 species worldwide and 55 in tropical America. Panama has just four species, and we cover two. Most Simaroubaceae have compound, alternate leaves, but the two described here are otherwise quite different in appearance. One interesting trait that unites the family is the presence of bitter chemicals called quasinoids, which have antimicrobial and antimalarial properties and are not known in other plants. Fresh leaves of Simaroubaceae have a bitter flavor if chewed as a result of these quasinoids.

Quassia amara (guabito amargo, cruceta, hom-

bre grande). A small tree of the forest understory. The

stem is typically leaning and there are often many separate stems on a single plant. Leaves are alternate, odd-compound, with three to five leaflets. The rachis between the leaflets is conspicuously winged. The leaves and bark have an unpleasant bitter flavor if chewed, hence the local name. The flowers are bright pink tubes with a narrow mouth, somewhat wider at the base, and are conspicuous in the dark forest understory during the brief flowering times. Fruits are small, fleshy, in clusters linked by a red receptacle. Distribution: Widespread in both Caribbean and Pacific lowlands, except the driest zones. Only in forest understory, and easy to find in the Canal Area from Gamboa north. Recognition: One of the first plants learned at Barro Colorado I. because of the winged leaf rachis; only lianas (the genera Paullinia and Serjania [Sapindaceae]) have similar leaves. Inga (Fabaceae—Mimosoideae) and Sapindus (Sapindaceae) are trees with winged leaf rachises, but both lack the terminal leaflet. When flowering, Quassia is conspicuous, and flowers are often found on the ground. A chemical with insecticidal properties can be extracted by boiling any part of the plant, and Quassia is a favorite local medicine.

Simarouba amara

Simarouba amara

Simarouba amara (aceituno, olivo). A tall forest tree. The trunk is straight, cylindrical, and unbuttressed. The bark on larger trees is light gray with splotches of dark and light green; it looks like camouflaging. Leaves are long, odd-compound, with 9–20 leaflets; juveniles generally have longer leaves with more leaflets than adults. Leaflets are smooth, shiny green, and asymmetric, with the outer half wider than the inner half; secondary veins are barely visi-

ble. The base of the petiole is swollen and flattened vertically. Broken leaves produce a clear resin that has a bitter taste. Flowers are tiny, green to yellowish, in large, loose clusters. Fruits are olivelike, turning from green to black, hence the common names. Distribution: Widespread in lowlands, including dry, wet, and lower montane zones. Common in mature forest at Barro Colorado I. and Soberania. Recognition: The many asymmetric and smooth leaflets are easy to learn. Dipteryx oleifera (Fabaceae—Papilionoideae) might be confused at first, because its leaflets are asymmetric, but it is very different. The other genus in Panama, Simaba, has two species with similar long, compound leaves; one in fact is so similar to Simarouba that it is easy to confuse. The genus Picramnia used to be considered part of the Simaroubaceae, but it is now placed in its own family, Picramniaceae (see Appendix 3).

Simaroubaceae

Quassia amara

Simarouba amara

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Siparunaceae

Siparunaceae Siparuna pauciflora

(Formerly Monimiaceae)

Siparuna pauciflora

A small family of treelets to trees in tropical forests, with a single species in Africa and 60 in the Americas. Panama has nine species in a single genus. Here we cover just one and omit a description of general family characters.

Siparuna pauciflora (pasmo hediondo, pasmo,

limoncillo). A treelet of forest understory. The trunk has low branches and often leans. Small branches are flattened and squarish in cross section. Leaves are simple and opposite, with fine teeth and short pubescence, feeling like a soft towel. Crushed leaves, especially young ones, have a strong odor, somewhat citruslike, but musky; it is quite different from other plant odors. The flowers resemble tiny mushrooms

growing out of the trunk or branches. They are flat caps on short stalks, with a small opening on top, initially green but maturing yellow; they never open and there are no petals. Crushed flowers produce the same odor as the leaves but more intensely. The fruit is a capsule that opens into a lovely, star-shaped display including a white fleshy backdrop and tiny seeds with red arils. Distribution: Very widespread in moist to wet and lower montane forests, including secondary and mature forests. Many grow along the outer part of Pipeline Rd., immediately adjacent to the road. Recognition: Few species have toothed, opposite leaves. Cassipourea elliptica (Rhizophoraceae) and Mollinedia dariensis (Monimiaceae, not covered) might be confused with Siparuna, but both have inconspicuous teeth and odorless leaves without pubescence. See also Crossopetalum parviflorum (Celastraceae). Panama’s eight other Siparuna species all have opposite leaves and the same strong and distinctive odor; some are toothed, others not. They are all lowland wet-forest to lower montane species. Siparuna used to be in the Monimiaceae with Mollinedia (see Appendix 3).

Turpinia occidentalis

Staphyleaceae Turpinia occidentalis

A small family of trees and shrubs with about 60 species worldwide, most common in temperate North America, where it is called the bladdernut family. Seventeen species are found in Central America and the Andes. Panama has just one species, so we omit the family description.

Turpinia occidentalis (cedrillo, cedrillo de mon-

taña). A widespread, medium-sized forest tree. On big trees, the trunk can be ribbed, with small buttresses. Leaves are opposite, compound and odd-pin-

nate, with opposite, toothed leaflets. Flowers are small and white, in loose terminal clusters. Fruits are small, three-sectioned, in loose clusters, turning black or purple when mature. Distribution: Generally a lower montane species in w. Panama and Costa Rica, but it is also widespread in lowlands of the Canal Area, though uncommon. Recognition: Should be unmistakable given the rare combination of opposite and compound leaves, but in the mountains you must consider Sambucus (Adoxaceae), which we do not cover but which likewise has opposite, odd-pinnate, toothed leaves. The family Brunelliaceae also has opposite, pinnate leaves, but it is only seen in very high mountains (we do not cover it; see Appendix 3). Otherwise, only the Bignoniaceae routinely have opposite, compound leaves, but the commonest bignon trees have palmate (Tabebuia) or bipinnate (Jacaranda) leaves. Spathodea, the introduced African bignon, has opposite, pinnate leaves, but it is not in forest and Turpinia is not in towns. A few Fabaceae have opposite, odd-pinnate leaves, but without teeth.

Staphyleaceae

Siparuna pauciflora

Turpinia occidentalis

451

452

Ternstroemiaceae

Ternstroemiaceae Ternstroemia tepezapote

(or Pentaphylacaceae, Formerly Theaceae)

Ternstroemia tepezapote

A modest family of shrubs and small trees found in tropical and temperate climates. There are nearly 300 species worldwide, with 120 in tropical America. It is a poorly known group, with two different family names being applied now; at one time, they were considered part of the tea family, Theaceae. Panama has six species, and we cover only one and omit the general family description.

Ternstroemia tepezapote (manglillo, manglillo de botón, manzanillo de sabana). A poorly known

forest tree. Bark is gray. Leaves are lustrous, shiny green, simple, alternate, untoothed, and bunched at the ends of branches. Secondary veins are inconspicuous. Flowers are showy and white, held singly on long stalks. Fruits are large, spherical, and inside have seeds wrapped in a red aril. Distribution: We find it all around the Canal Area in lowland forest, but it is rare. Otherwise it is known mostly in mountains and some very wet lowland sites. Recognition: The combination of very bunched and very shiny leaves with inconspicuous veins and no latex sets it apart, but each of those characters shows up in other families. This species is rarely seen and so is easy to forget. The family contains three more genera: Cleyera (one species), Freziera (three), and Symplococarpon (one). All are montane, small to medium-sized trees with large flowers. Freziera is easiest to recognize, having shiny green leaves with brown undersides.

Daphnopsis americana

Thymelaeaceae Daphnopsis americana

A modest family of about 400 species worldwide, but having only 75 species in tropical America. Most are shrubs or small trees, but there are herbs, lianas, and large trees as well. Only three species are found in Panama, and we describe one. The family generally has simple, alternate leaves, and the three Panamanian species all do. The most distinctive character is very tough bark, and tearing a leaf off brings a long string of bark off the branchlet. Otherwise, the family is not well characterized.

Daphnopsis americana. A rare, medium-sized tree. The bark is gray. Twigs have white lenticels.

Leaves are simple, alternate, fairly small, smooth, and are somewhat bunched toward the ends of branches, held at various angles. Flowers are small, yellow, held in dense, round-topped heads. Fruits are berries that are snow-white when mature. Distribution: Mostly a cloud-forest species through Panama and Costa Rica, with some records in lowland wet forests. Recognition: A nondescript species, not often seen and thus difficult to learn. Similar small, clustered leaves in other species are not no smooth. If you think of tearing a leaf off, the tough bark can be helpful. Another species in the genus, D. correae, has larger leaves that are not as smooth and it is more likely to be confused with other families than with D. americana. Panama has one more genus in the family, Schoenobiblius, with one species. It has simple, alternate leaves, and is sparsely distributed at a few wet and montane sites.

Thymelaeaceae

Ternstroemia tepezapote

Daphnopsis americana

453

454

Ulmaceae

Ulmaceae

Ulmus mexicana

Ulmus mexicana

The elms comprise a small but well-known temperate family. Only about 35 species are known worldwide, with 13 in the American tropics. We cover here just one of the four species in Panama and omit a family description.

Ulmus mexicana (cenizo, elm). A tall tree of highmountain forest. Big trunks can be swollen into mod-

erate buttresses. The bark is gray, fissured, with small pieces peeling free. Leaves are simple, alternate, toothed, asymmetric at the base, and rough to the touch. Flowers are tiny and white. Seeds are winged and have a fringe of small hairs on the edge. Distribution: Only above 2000 m elevation in far w. Panama through Costa Rica. Recognition: If you know elms in North America or Europe, you will recognize the asymmetric base and rough leaves. Guazuma ulmifolia (Malvaceae—Byttnerioideae) has somewhat similar leaves, but does not occur at high elevations. The other genus in Panama, Ampelocera, has three species in the lowlands. They have asymmetric leaf bases, but lack teeth. Trema and Celtis (now Cannabaceae) used to be classified within Ulmaceae.

Cecropia garciae

Urticaceae Cecropia garciae

A major family of about 1300 species worldwide, but most are herbs or small, soft shrubs, including the stinging nettle of Europe and North America. There are about 450 species in the New World tropics, but most are herbs. The Panama checklist has 24 trees and treelets. We illustrate five species and describe

one more. These tree Urticaceae have at one time or another been classified either as Cecropiaceae or Moraceae. The species we cover are all highly distinctive trees. The biggest genus, Cecropia, includes the most important pioneer trees in the Americas, commonly invading nearly every roadside and forest clearing. They are among the fastest-growing trees in the world. Moreover, they are among the most obvious and easily recognizable trees anywhere, with huge, umbrella-shaped leaves confined to the top of the tree, and trunks covered with leaf scars. Within Cecropia, though, species can be difficult to tell apart. The other genus of trees, Pourouma, has similar leaves and is also easy to recognize.

Cecropia insignis

Cecropia insignis

they have fallen, but look mostly greenish while still on the tree. In C. garciae, they really look white.

Cecropia insignis (guarumo). A tall tree of mature

Cecropia garciae. Not illustrated. This species is alphabetically first, but the full descriptions come later for the more widespread species. C. garciae is like all the other Cecropia in general form, but leaf undersides are very white and deeply lobed, so each lobe is almost separate from the next. Distribution: Only known in wet and montane forest of c. Panama. Found at Pipeline Rd. Recognition: Leaves of other Cecropia species look white on the underside after

forest clearings. The trunk is tall, light colored, cylindrical but often leaning, and has circular scars throughout where old leaves have fallen. Trees of moderate size or larger have stilt-roots at the base. The huge leaves are only near the top of the tree and are like umbrellas, with the petiole attached at the center. Each leaf has six to nine broad lobes that are rounded at their tips; there is a scaly feel to the upper side, and the underside has a reddish tint, especially on the veins, due to fine hairs. Flowers are minute, white, held on spikes resembling a set of fingers dangling downward just beneath the leaves. These same fingers develop into green, fleshy fruits. Distribution: Widespread in lowlands, from dry to wet zones. Unlike other Cecropia, nearly always found in mature

Urticaceae

455

Ulmus mexicana

Cecropia insignis forest, not open areas or forest edge. Inside the forest it is always in the sun; juveniles are only found in natural forest clearings, whereas tall adults are widely scattered in the canopy. Recognition: The genus Cecropia cannot be confused. No other tree has umbrella-like leaves, with the petiole attached in the leaf

center, and the trunks with ring scars are also distinctive. Distinguishing species of Cecropia, however, is not so easy. C. insignis is most like C. peltata, with moderately deep leaf lobes. Check for reddish hairs under the leaves, which distinguish C. insignis.

456

Urticaceae

Cecropia longipes

Cecropia longipes

Cecropia longipes (guarumo). See the description of C. insignis, which this closely resembles in overall trunk and leaf form. C. longipes, however, has only slightly lobed leaves, so the leaf is almost round. Flowers and fruits differ too, being held on very long stalks. Distribution: Well known in the Canal Area, but otherwise sparse and not seen in Costa Rica. Like the entire genus, restricted to sunny areas, and found in mature forest gaps as well as edges and open areas. Recognition: Quite similar to C. insignis and C. peltata, but shallowly lobed leaves should distinguish it, and big trees often have the long-stalked fruits.

Cecropia obtusifolia

Cecropia obtusifolia (guarumo). A tall forest tree of clearings and edges. Just like other Cecropia in most respects, but the leaves have more numerous and narrower lobes. Distribution: Widespread in lowlands and mountains, dry to wet. This is one of the common Cecropia invading clearings and edges throughout Panama. Recognition: This is the easiest Cecropia to identify, because the leaves have narrow lobes.

Cecropia obtusifolia

Urticaceae

Cecropia longipes

Cecropia obtusifolia

457

458

Urticaceae

Cecropia peltata

Cecropia peltata

Cecropia peltata (guarumo). See C. insignis.

Distribution: Known especially along the Pacific coast, including the driest regions, but also shows up at wet and lower montane sites. One of the common

Cecropia invading clearings and edges. Seldom found in mature forest clearings; nearly always in secondary forests or open areas. Recognition: The leaves closely resemble those of C. insignis, with moderately deep lobes, but they lack the reddish hairs. The two species seldom grow together, because C. insignis is in gaps inside the forest while C. peltata is nearly always outside the forest. Five more Cecropia are found in Panama. C heterochroma is a wet forest tree of c. and w. Panama with young leaves that are bright red below. The other species are rare in Panama.

Pourouma bicolor

Pourouma bicolor

Pourouma bicolor (uvito, mangabé, guarumo macho). A forest tree with umbrella leaves. The trunk is very similar to that of Cecropia, with rings throughout the length and stilt-roots at the base. As in Cecropia, leaves are large, with wide lobes, whitish on the underside, but with important distinctions. First, the petiole does not attach in the center of the leaf, as in Cecropia, but at the base (as in most plants). Second, leaf lobes of Pourouma are triangular, pointed at the tip, not rounded as in most Cecropia. Flowers are small, reddish brown, in clusters below leaves that are borne on reddish stalks. Fruits are large berries. Distribution: In lowland forest of moist to wet zones, sometimes lower montane. Largely confined to forest openings, but does not invade edges and clearings like Cecropia, and generally not nearly as abundant as Cecropia. Recognition: With a little care, this species is easy to be certain about. From the trunk and large leaves, it can be confused with

Cecropia, but look carefully at the petiole joining the leaf. All these species drop lots of leaves on the ground, so it is easy to check this. Pourouma can also be confused with Sterculia apetala (Malvaceae—Sterculioideae), which has leaves of the same shape. Usually leaves of Sterculia are smaller, but in juvenile trees they might match the size of those of Pourouma. Leaves of Pourouma are whitish on the underside, though, like those of Cecropia but very unlike those of Sterculia. Panama has two other Pourouma species, both rare. One has lobed leaves, but the other has small unlobed leaves. Both have white undersides like those of Cecropia and P. bicolor. Urticaceae includes four more tree genera, but none are at all like Cecropia. Boehmeria has herbs or shrubs of montane areas, with two species just qualifying as small, shrublike trees. Pouzolzia has two similar species occurring widely at low elevation. The other two genera include some commonly seen treelets. Myriocarpa longipes is a widespread species, from lowland to lower montane and dry to wet, of streamsides and forest edge. It is abundant along streams at Pipeline Rd., easily recognizable from big, rough, heart-shaped leaves. The genus Urera includes six species with similar heart-shaped leaves. Be careful; Urera, like the nettles of North America, has stinging hairs on leaves, petioles, and flowers.

Urticaceae

Cecropia peltata

Pourouma bicolor

459

460

Verbenaceae Citharexylum caudatum

Verbenaceae Citharexylum caudatum

A large family of about 1000 species worldwide, found in nearly every habitat from temperate to tropical, and including trees, shrubs, herbs, and lianas. There are approximately 300 species in the American tropics. They are closely related to the mint family, Lamiaceae, and several important trees formerly classified within the Verbenaceae are now Lamiaceae. Panama has 18 species of the remaining Verbenaceae, of which we cover 1. Verbenaceae have opposite leaves, and the stems are often square or angled, both traits shared with the Lamiaceae. Most Verbenaceae have toothed leaves, but the biggest genus and the one we illustrate does not. Flowers are characteristic: small tubes with flattened and slightly asymmetric lips at the opening. If you know mint flowers, they are similar.

Citharexylum caudatum (manglillo, moco de pavo, palomo). A medium-sized tree of roadsides and

parks. Leaves are simple, opposite, untoothed, smooth, and spaced regularly along branches. At the base of each leaf are tiny raised glands, one on either side of the main vein. Flowers are white, borne on terminal spikes. Fruits are orange, fleshy, on the same spikes. Distribution: Widespread but sporadic. Most often seen in Caribbean coast lowlands, but also fair commonly in and around Panama City and sometimes in lower mountains. Usually seen along roads or in other open areas. Recognition: Looks a lot like a mangrove, especially Laguncularia (Combretaceae), which even has similar glands. But Laguncularia has smoother leaves with weak venation. Citharexylum might be confused with Myrtaceae, but most Myrtaceae have a collecting vein. Eight more Citharexylum species are found in Panama. Several are montane, and others very rarely seen. They all have opposite leaves, but some Citharexylum have teeth. Verbenaceae includes two more genera in the region, Duranta (three species) and Lippia (five). Both are montane but rare. Duranta is a spiny shrub and Lippia is a small tree. The shrubby Lantana—a garden bush and weedy pest in much of the world— is in this family and native to Panama, but we do not count it as a tree.

Hybanthus prunifolius

Violaceae

Hybanthus prunifolius

This is the violet family, a moderately large group found from the lowland tropics to temperate climates, having 800 species worldwide. It is best known for the large genus Viola, a garden plant and familiar forest-floor herb of North America and Europe, but other Violaceae are widespread in the tropics, and many are woody shrubs, lianas, or small trees. In tropical America, there are 200 of these woody forest species, and another 100 species of Viola in the mountains. The Panama checklist has 24 species of treelets or small trees, and we cover 2 very common ones. Indeed, although not especially rich in species, many tropical forests throughout the world have abundant treelets of this family in the understory. The family is diverse in form and difficult to characterize. The standard Violaceae have alternate, toothed leaves spaced regularly along branches, like

Salicaceae. But only one of the common genera we cover follows this pattern. The other has opposite, weakly toothed leaves, and other genera (not covered) have no teeth at all. Some of the tropical treelets have violet-like flowers, with an expanded lower petal, but most have simple white flowers that are either symmetrical or have one slightly expanded petal. You will probably learn to recognize the two common genera on their own characters before you learn the tropical Violaceae as a group.

Hybanthus prunifolius (flores de lluvia). A treelet of the shaded forest understory. The stem typically leans, especially in larger plants, and has irregular branches. Leaves are alternate, toothed, with a long, pointed tip, and are spaced regularly along branches. Stipules are whitish and persist well, so some are always present. Flowers are fairly large, mostly white but with yellow marks on the enlarged lower petal, produced after rains during the dry season. Fruits are green capsules. Distribution: This is the dominant understory plant of old-growth forest of Barro Colorado I., so abundant that there is nearly always a Hybanthus within sight. Fairly common in Soberania, but otherwise seldom seen. Recognition: The white stipules are the key, because other species with similar leaves have nothing like them.

Violaceae

461

Citharexylum caudatum

Hybanthus prunifolius

The most similar species is Turnera panamensis (family Turneraceae, not illustrated; see Appendix 3), the toothed leaves of which are the same size and shape as those of Hybanthus, but Turnera lacks the whitish stipule. Several Salicaceae, especially the genus Casearia, have toothed leaves like those of Hybanthus, but usually are larger trees. Hyban-

thus is quite a sight in the dark understory when it flowers. Two more Hybanthus species are found in Panama, both shrub-sized with toothed, alternate leaves and whitish stipules. Both are rarely seen anywhere; outside the Canal Area, all three species are rare.

462

Vochysiaceae Rinorea sylvatica

Rinorea sylvatica

Rinorea sylvatica. A treelet of forest understory.

Leaves are simple, opposite, and tend to be bunched toward the branch tip; there can be a third leaf joining the opposing pair at the end of the branch. The leaves are toothed, but sometimes inconspicuously so. The leaf base is asymmetric, sometimes clearly and sometimes subtly, and the base almost folds against the branch. Flowers are tiny, yellow, on long terminal stalks. Fruits are little green capsules. Distribution: Commonly seen at Barro Colorado I.; otherwise widespread but seldom seen. Only in forest shade. It grows in dense clusters between which

there are none at all. Recognition: Traits of Rinorea are variable and it might seem like a difficult plant to learn. But its abundance around Barro Colorado I. makes it easy there, where it will eventually be learned from the opposite, bunched leaves and the terminal pair that (often) has leaf bases folded toward the branch. Ten more Rinorea species are found in Panama. They can usually be known by opposite, slightly toothed leaves bunched toward the branch tip, sometimes with a third leaf at the tip. Several are common in mature forest understory. Violaceae has four more tree genera in Panama: Amphirrhox (one species), Gloeospermum (six), Leonia (one), and Paypayrola (two). None have teeth, so they will not be confused with Rinorea or Hybanthus. All have alternate leaves and may be confused with many other families. None are seen very often, though Gloeospermum and Leonia show up regularly in wet Caribbean forest.

Vochysiaceae Vochysia ferruginea

Vochysia ferruginea

A small family of about 200 tropical tree species, nearly all in the Americas (there are three species in Africa). Panama has nine species, and we cover the single very abundant one. Vochysiaceae are similar and closely related to Myrtaceae. Both families have opposite leaves that tend to be regularly spaced along branches. Myrtaceae is one of the three dominant opposite-leaved families (with Rubiaceae and Melastomataceae), and are usually recognized by a marginal collecting vein. Some Vochysiaceae have this marginal vein, but in others it is obscure. There are many more Myrtaceae than Vochysiaceae, and you are probably better off thinking Myrtaceae for an opposite-leaved tree with collecting veins and learning the individual Vochysiaceae.

Vochysia ferruginea (mayo, flor de mayo, botarrama, tecla). A large, abundant tree of secondary forest. The trunk is straight and cylindrical, and the crown has a flat top, with all branches reaching

about the same height. Bark is dark with white or gray spots. Twigs are square and have fine red hairs. Leaves are simple, opposite, narrow and pointed, and veins below are reddish with the same fine hairs. Flowers are bright yellow, densely held on spikes, and big trees are covered in these spikes at the top of the crown. The species flowers for weeks in April and May and again (less so) in September. Fruits are dry capsules that turn dark and split open, releasing tiny winged seeds. Distribution: Widespread in lowlands everywhere. One of the most abundant trees in secondary forests and wooded roadsides around the Canal; much less common in old growth. Recognition: Its abundance makes it easy to learn. The reddish veins on opposite leaves are distinctive. Vochysia is very easy to pick out on distant hillsides when flowering. Vochysia includes four more species. V. guatemalensis is widely distributed in wet lowlands and lower mountains of Costa Rica, but rare in Panama. It has larger leaves than those of V. ferruginea, held in groups of three. Another species is restricted to c. Panama, and the last two are rare. Two more genera of the family, Qualea (two species) and Erisma (one), are known in Panama. One Qualea is quite common in wet forests near the Canal, and has easily recognized, narrow, shiny, opposite leaves with glands at their base. Erisma was only recently discovered in Panama in our inventories. It also has opposite leaves.

Vochysiaceae

Rinorea sylvatica

Vochysia ferruginea

463

Part III

Supporting Material

Appendix 1

Terminology Some botanical terminology may be unnecessary jargon, but there are precise terms for common characters that make it much easier to write about tree species. Here are those we use regularly in the text. There is an entire book on terminology (Harris and Harris 1994) and a great Wikipedia Glossary of Botanical Terms.1 Alternate. Describes leaves (or leaflets) that grow singly off a branch (or rachis), that is, not in pairs or groups; see drawing in Figure 2.1 (Chapter 2). Branches off a trunk can also be described as alternate. Bifoliate. Describes a compound leaf with exactly two leaflets. Bipinnate. Describes a compound leaf with two divisions: the leaf is divided into leaflets, and the leaflets are divided into smaller units called pinnules (a good descriptive term would be “subleaflets,” but botanists do not use it). There are also tripinnate leaves, with a third division, but these are rare. Digitate. Describes compound leaves and means the same as palmate. Distichous. Describes leaves that grow on either side of a branch, held in a flat plane, and regularly spaced. Even-pinnate. Describes a pinnate leaf in which there is no terminal leaflet, most often used where all leaflets are in opposite pairs. The botanical jargon is paripinnate. We prefer the easier-to-remember “even-pinnate.” The opposite condition is odd-pinnate or imparipinnate. Imparipinnate. Difficult-to-remember botanical jargon for a pinnate leaf with an odd number of leaflets. See odd-pinnate, which we prefer, and also paripinnate and even-pinnate. Interpetiolar stipule. A stipule that is attached to the branch between the two leaf petioles; leaves must be opposite to have it. When such a stipule falls, the scar left be1

http://en.wikipedia.org/wiki/glossary_of_botanical

_terms.

hind is a tiny ridge crossing the branch from one petiole to the other. Leaflet. In a compound leaf, the single unit of leaf tissue. In most cases, a leaflet looks just like a leaf. See drawings in Figures 2.2–2.4 (Chapter 2). Lenticel. A tiny pore on branches or trunks, sometimes raised or swollen like a wart. Odd-pinnate. Describes a pinnate leaf in which there is a single leaflet at the end (as well as additional leaflets arranged along the rachis). It is most often used where the remaining leaflets are in opposite pairs, so that the terminal leaflet is conspicuous, and there is an odd number of leaflets. The botanical term is imparipinnate. The opposite condition is even-pinnate or paripinnate. Opposite. Describes leaves (or leaflets) that come in pairs, with both members of a pair growing from the same spot on a branch (or rachis), that is, not singly; see drawing in Figure 2.1 (Chapter 2). Branches off a trunk can also be described as opposite, and species with opposite leaves generally also have opposite branches. Palmate. Describes a compound leaf in which the leaflets are arranged like spokes of a wheel, all arising from a single point. Paripinnate. A difficult-to-remember botanical term for pinnate leaves with an even number of leaflets. See even-pinnate, which we prefer, and also imparipinnate and odd-pinnate. Petiole. The stalk holding the leaf.“Leafstalk” would be easier to remember, but petiole is so widely used it is best to know. Pinna. A leaflet. This is the seldom-used technical term for the units of leaf tissue in com-

468

Appendix 1

pound leaves. Plural is pinnae. Even botanists use “leaflet” over “pinna,” but see pinnule. Pinnate. Describes a compound leaf in which leaflets are arranged on either side of a single rachis. This is by far the most common type of compound leaf. Pinnule. The units of leaf tissue in bipinnate leaves, essentially a “subleaflet,” but no one uses that word, so this technical term is necessary. Pubescence. Small hairs. There are a variety of words describing precise kinds of hairs, but the differences can be distinguished only with magnification, so we use this one word to describe many kinds of hairs found on twigs, petioles, or leaves. There is nothing especially wrong about calling a leaf “hairy,” but pubescence is typically too small to see with the naked eye, unlike the hair of mammals, so in this case the technical term is more precise. Pulvinus. The swollen base of a petiole. Rachis. The stalk holding the leaflets of a pinnately compound leaf. There is no good common word for this. Stipule. A tiny flap of tissue at the base of a leaf, just where the petiole meets the branch.

It can be green, resembling a tiny leaf, or brown, like a dried leaf. Sometimes it is narrow and pointed, barely more than a little filament. In many cases, the stipule falls off when a leaf is still young; in some cases, a tiny ridge is left behind as a scar. There is no common word for stipules. Subopposite. Describes leaves that arise almost but not quite precisely opposite. Trifoliate. Describes a compound leaf with exactly three leaflets. Tripinnate. See under bipinnate. Two-ranked. A nontechnical term for distichous, defined previously. Verticillate. Describes several branches emerging from a single point on the trunk. Two branches emerging from opposite sides of the trunk are called opposite. See whorled, which is essentially the same, but botanists generally call branches verticillate and leaves whorled. Whorled. The condition in which three or more leaves grow from a single spot on a branch. Whorled leaves are neither alternate nor opposite. Whorled can be applied to branches, and thus would be the same as verticillate, but whorled usually refers to leaves.

Appendix 2

Major Leaf Traits of Tree Families Known in Panama and Costa Rica A tabulation of the basic leaf form of tree families known in Panama and Costa Rica is presented here. Of 134 native families, 117 are included. The 17 excluded either barely qualify as trees, or are very easily recognized: Apiaceae, Araceae, Asteraceae, Cactaceae, Connaraceae, Cyatheaceae, Ericaceae, Gentianaceae, Gesneriaceae, Liliaceae, Loranthaceae, Menispermaceae, Onagraceae, Passifloraceae, Poaceae, Polygalaceae, and Zamiaceae. Brief descriptions of those 17 families appear in Appendix 3. Although built around Panama’s tree species and families, this table is just as applicable in Costa Rica. The traits apply equally well in both countries, and 131 of the 134 native tree families from Panama also occur in Costa Rica (Buxaceae, Cyrillaceae, and Goupiaceae do not; we have not, however, searched for families known in Costa Rica but not in Panama). In most cases, the family traits hold elsewhere in the world, and this table should work well in South America. In Africa and Asia, though, there are more likely to be exceptions and this table is not designed for those regions. Families are divided into four groups based on leaf form and arrangement: alternate-simple, alternate-compound, opposite-simple, and opposite-compound (as in Table 2.1 in the text). Fourteen families appear twice because they fall in two categories: seven families that include some species with alternate leaves and others with opposite leaves are marked AO, and

seven that include some species with simple and others with compound leaves are marked SC. The columns “country” and “book” give the number of species known in all of Panama and the number described in this book, respectively. The 25 families appearing in Table 2.1 are noted as “major”; these are the most important families, found in most forests of the region. Twentyseven families are marked “scarce”; those are rare and seldom encountered. The remaining unmarked families are somewhere between “major” and “scarce”; they are fairly widespread but not common and absent at many sites. Other leaf characters are indicated by an “x” in the appropriate column. Families are marked “x” if many species in a family possess the trait, but this does not mean that all species in a family have the trait! In particular, teeth and clustered leaves are often inconsistent within families. This table is intended mainly to indicate which families are plausible given a set of observations about leaves on a tree. Good sources for more thorough descriptions of family characters are Gentry (1996) and Smith et al. (2004). Besides the major leaf traits, there is also an indication “montane.” This column denotes families restricted to lower or upper montane forest, never seen in lowlands. The remaining families occur widely from lower montane into lowland zones.

71

7

Euphorbiaceae

16

Elaeocarpaceae

Erythropalaceae

8

6

Ebenaceae

2

1

Dilleniaceae

Escalloniaceae

1

4

Dichapetalaceae

Erythroxylaceae

1

1

Cyrillaceae

1

14

0

3

2

1

0

1

6

2

1

8

3

Clethraceae

9

Combretaceae

32

Chrysobalanaceae

1

2

0

5

2

10

1

1

3

0

16

28

8

0

1

Book

Cochlospermaceae

26

8

Caricaceae (SC)

Celastraceae (AO)

2

Cardiopteridaceae

2

Bixaceae

21

1

Betulaceae

Capparaceae

47

Araliaceae (SC)

4

8

Aquifoliaceae

29

36

Apocynaceae (AO)

Cannabaceae

85

Annonaceae

Boraginaceae

5

12

Anacardiaceae (SC)

simple

5

Country

Actinidiaceae

Family

Achariaceae

Leaf

Alternate-

major

scarce

scarce

major

scarce

scarce

major

scarce

major

major

major

Importance

x

x

Lobed

Table A.1 Major Leaf Traits of Tree Families Known in Panama and Costa Rica

x

x

x

x

x

x

x

x

x

x

x

x

x

Toothed

x

x

x

x

x

Latex

x

x

x

x

x

x

Two-ranked

x

x

x

x

x

x

x

x

x

x

x

Clustered

x

x

x

x

x

x

x

x

x

x

x

x

Stipule

x

x

x

Montane

2

7

4

Putranjivaceae

1

1

4

Proteaceae (SC)

1

4

24

Polygonaceae

1

0

4

1

1

0

1

2

0

6

9

1

1

15

0

22

1

Podocarpaceae

41

1

Piperaceae

15

Phyllonomaceae

1

Opiliaceae

Phyllanthaceae

2

Olacaceae

1

12

Ochnaceae

1

1

Papaveraceae

86

Myrsinaceae

Nyssaceae

Pellicieraceae

1

21

Myristicaceae

2

Muntingiaceae

Myricaceae

1

85

97

Malvaceae (SC)

Moraceae

5

Magnoliaceae

Metteniusaceae

6

1

1

39

Lecythidaceae

Lepidobotryaceae

2

1

3

9

3

Lacistemataceae

1

0

134

3

Icacinaceae

Lauraceae

4

5

Hernandiaceae

Humiriaceae

9

1

Fagaceae

Goupiaceae

Alternate-

simple

scarce

scarce

scarce

scarce

major

major

scarce

major

major

major

scarce

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

0

60

1

4

7

6

Solanaceae

Stemonuraceae

Styracaceae

Symplocaceae

Ternstroemiaceae

17

Burseraceae

Juglandaceae (AO)

Fabaceae

2

262

4

50

Arecaceae

Caryocaraceae (AO)

12

47

Anacardiaceae (SC)

Araliaceae (SC)

Alternate-

compound

8

26

Violaceae (AO)

Caricaceae (SC)

5

24

Turneraceae

Urticaceae

2

Ticodendraceae

Ulmaceae

3

1

Thymelaeaceae

2

2

Schoepfiaceae

5

80

Sapotaceae

Theaceae

43

Salicaceae

Theophrastaceae

1

12

Sabiaceae

0

0

75

0

2

6

11

3

8

2

7

1

0

0

1

0

0

0

0

0

0

13

12

0

5

5

24

3

Book

Rosaceae (AO)

simple

10

Country

Rutaceae (SC)

Family

Rhamnaceae (AO)

Leaf

Alternate-

Table A.1 (Continued)

scarce

major

scarce

scarce

major

major

major

major

major

scarce

scarce

scarce

scarce

major

major

scarce

Importance

x

x

x

Lobed

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

Toothed

x

x

x

Latex

x

x

x

x

x

Two-ranked

x

x

x

x

x

x

x

x

x

x

x

Clustered

x

x

x

x

x

x

x

x

x

x

Stipule

x

x

x

x

x

x

x

Montane

97

26

7

Celastraceae (AO)

Chloranthaceae

5

21

5

2

25

Hypericaceae

Lamiaceae

Loganiaceae

Lythraceae

Malpighiaceae

2

3

10

Nyctaginaceae

Oleaceae

Quiinaceae

Rhamnaceae (AO)

5

15

Myrtaceae

Rhizophoraceae

5

69

Monimiaceae

152

1

1

Cornaceae

Garryaceae

Melastomataceae

3

1

0

16

major

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

2

x

x x

x

x

x

x

x

x

x

x

x

x

3

x

x

scarce

major

major

scarce

scarce

major

scarce

major

scarce

major

major

0

0

2

8

0

17

5

1

0

4

3

0

1

1

Buxaceae

0

1

3

2

11

5

1

0

14

36

Apocynaceae (AO)

22

11

47

1

Alzateaceae

Clusiaceae

4

Adoxaceae (SC)

4

Simaroubaceae

simple

35

Sapindaceae

11

24

Rutaceae (SC)

Acanthaceae

7

Opposite-

6

Proteaceae (SC)

36

Picramniaceae

Malvaceae (SC)

Meliaceae

Alternate-

compound

x

x

x

x

x

Rubiaceae

simple

1

Staphyleaceae

1

0

0

5

2

Cunoniaceae

0

0

3

4

Brunelliaceae

Caryocaraceae (AO)

Juglandaceae (AO)

7

1

0

1

2

1

1

1

32

0

Book

24

Bignoniaceae

1

Winteraceae

compound

10

Vochysiaceae

4

26

Violaceae (AO)

Adoxaceae (SC)

18

Verbenaceae

Opposite-

11

4

248

5

Country

Siparunaceae

Scrophulariaceae

Family

Rosaceae (AO)

Leaf

Opposite-

Table A.1 (Continued)

scarce

scarce

scarce

scarce

major

scarce

major

scarce

major

scarce

Importance

Lobed

x

x

x

x

x

x

x

x

x

x

Toothed

Latex

x

Two-ranked

x

x

Clustered

x

x

x

x

x

x

x

Stipule

x

x

x

x

x

x

x

Montane

Appendix 3

Families Not Included Brief descriptions are given of 51 plant families with trees or treelets in Panama that are not covered in the main text. Three additional names are included because they were once commonly used but are no longer valid in Panama. The families listed here are considered in the introductory guide to family identification (Chapter 2) and included in the table of leaf characters (Appendix 2). Actinidiaceae. A single genus, Saurauia, of four montane plus one lowland tree species, the latter known at outer Pipeline Rd. and other areas on the Caribbean side of the Canal. They have alternate, toothed leaves clustered at the ends of branches. Alzateaceae. One species of small or mediumsized tree in the genus Alzatea, from montane forests. It has stilt-roots and thick, round, opposite leaves. Apiaceae. The carrot and parsley family has two species of the genus Myrrhidendron that can be a few meters tall. They occur near the tops of the highest mountains. Aquifoliaceae. Eight species of the holly genus, Ilex, are found in montane forest, with one also widespread in Caribbean lowlands. They are medium-sized trees with nondescript, simple, alternate, and usually toothed leaves. Araceae. The Philodendron family, a group of large-leaved herbs that often grow on trees and are a quintessential tropical forest component, has one species of Montrichardia that can have a woody stem 2–3 m tall. Brunelliaceae. Three montane species of Brunnellia are small or medium-sized trees with opposite, pinnately compound leaves and toothed leaflets, a very rare combination of characters most commonly found in Bignoniaceae. Buxaceae. The boxwood family, genus Buxus, has a single small tree species known from just a handful of specimens in c. Panama. It has opposite, shiny green leaves with a collecting vein parallel to the margin. Cactaceae. This well-known family has one species of Pereskia with a woody stem, sharp

spines, and leaves; it is nothing like a typical cactus. Acanthocereus and Opuntia, which are classic columnar cacti, tall enough to be considered trees, each have a single species in Panama. Cardiopteridaceae. Two genera, Citronella and Dendrobangia, both formerly in the Icacinaceae (which is covered in the text), have a single large tree species each, the former in high mountains, the latter in lowland wet forest. Leaves are simple, alternate; flowers tiny and white. Easy to confuse with several other families. Caryocaraceae. This small Neotropical family has one species of Anthodiscus and three of Caryocar, all tall trees confined to very wet sites and not common. All have toothed, trifoliate leaves. Connaraceae. An important liana group in lowland tropics, with a few treelets, including three seldom-seen species in Panama in the genera Connarus and Rourea. They have pinnately compound leaves and petioles that look just like those of Fabaceae (and are easily confused). Cunoniaceae. An important high-mountain group of medium-sized trees with toothed, opposite leaves, usually pinnately compound. Five species in the single genus Weinmannia are known in Panama; the group is more widespread and better known in Costa Rica. Cyatheaceae. The tree-fern family has five species in the genera Alsophila and Cyathea. They look just like ferns on tall woody stems, and some are common and easy to spot in wet or montane forest. Dichapetalaceae. Trees or treelets (plus lia-

476

Appendix 3

nas) with simple, alternate, untoothed leaves with no easy distinctions. The genera Dicha­ petalum, Stephanopodium, and Tapura are known as trees, with four species total; Dicha­ petalum is seen most often, with one species widespread in lowland wet forests and another montane. Ericaceae. Most are woody epiphytes in the tropics, but Bejaria, Comarostaphylis, and Vaccinium include four treelet species in montane forests. The first genus resembles Rhododendron; the last is the blueberry genus. Escalloniaceae. Escallonia has two treelet species in very high mountain forest. They have simple, alternate, finely toothed leaves. This genus was formerly in the Saxifragaceae. Flacourtiaceae. Defunct. Most moved to Salicaceae and some to Achariaceae, both of which are covered in the text. Garryaceae. A single species of medium-sized tree in the genus Garrya is found in high mountains of far w. Panama and Costa Rica. Leaves are simple, opposite; flowers and fruits are hanging tassels. Gentianaceae. The gentian family has two tree genera, Macrocarpaea and Potalia, with one species each, the former montane, the latter in Caribbean lowlands. They are small treelets with opposite leaves, otherwise quite different from one another. Gesneriaceae. Most are soft herbs, and many are epiphytes, but one species each of Besle­ ria and Peltanthera are medium-sized trees with opposite leaves of mountains or wet lowlands. Goupiaceae. One species of Goupia is a medium-sized tree with alternate, toothed leaves on long branches, known only in e. Panama lowlands. It was formerly in the Celastraceae. Hippocastanaceae. Defunct. Moved to Sapindaceae. It used to include the genus Billia in Panama (mentioned in the text but not illustrated). Juglandaceae. The walnut family is mostly north temperate, but the genera Alfaroa and Oreomunnea include a single tall tree species each in high mountains of Panama and

Costa Rica. They have compound leaves with many toothed leaflets. Liliaceae. Lilies are the familiar large-flowered herbs, but there are a few tree species. A single species of the genus Dracaena is found on the Pacific slope. It has long, thin, bladelike leaves on an erect stem, like the Joshua tree (Yucca) of the sw. United States. Loganiaceae. The genus Strychnos has four species growing as small trees, plus several more lianas. They have simple, opposite leaves with three major veins (see Caryo­ daphnopsis in the Lauraceae). Loganiaceae formerly included Potalia (now Gentianaceae) and Peltanthera (now Gesneriaceae). Loranthaceae. This is the mistletoe family, but a single species in the genus Gaiadendron is a free-standing tree, though short and multistemmed, found at high elevation. Menispermaceae. Most are lianas, but the genus Hyperbaena has two medium-sized trees. They have simple, alternate leaves with thickened petiole bases. Metteniusaceae. Just one rare tree species in the genus Metteniusa is known in Panama. It has simple, alternate leaves, and used to be classified in the Icacinaceae (which is covered in the text). Monimiaceae. The genus Mollinedia includes five tree species; some are fairly common and widespread. They have opposite leaves with teeth. This family used to include Sipa­ runa, now in the Siparunaceae and covered in the text. Nyssaceae. A single species of tall trees in the genus Nyssa is found in high mountains of w. Panama and Costa Rica. It has simple, alternate leaves. Olacaceae. The genus Ximenia has a single species of small, spiny trees with simple, alternate leaves. This family also used to include Heisteria, now in the Erythropalaceae and covered in the main text. Oleaceae. The olive family has one species each in the genera Chionanthus and Forest­ iera, mainly in high mountains. They have simple, opposite leaves. Onagraceae. The fuchsia family has one treelet species in the genus Fuchsia and another

Families Not Included

in the genus Ludwigia. Both are restricted to high mountains. Opiliaceae. A single species of Agonandra is a tall tree of dry areas, with simple, alternate, small leaves. Passifloraceae. The passion-flower genus, Pas­ siflora, has close to 50 species, all but one of which are vines. The one is a small tree of high mountains. Phyllonomaceae. One treelet species in the genus Phyllonoma, formerly in the Saxifragaceae, occurs in the mountains of w. Panama and Costa Rica. It has simple, alternate, toothed leaves. Bizarrely, flowers are born on the leaf blade, toward the tip. Picramniaceae. The single genus, Picramnia, has five species. Picramnia has alternate, compound leaves very like those of Fabaceae, and we mention P. latifolia in the text under Pterocarpus rohrii of the Fabaceae (Papilionoideae), which it greatly resembles. Picramnia was formerly in the Simaroubaceae, which is covered in the main text. Poaceae. The grass family includes bamboos, which qualify as trees. One native species in the genus Guadua is found in Panama, plus many species introduced from Asia or elsewhere in South America. Polygalaceae. Mostly lianas or herbs, but one species of Polygala is a rare tree. It has opposite leaves with petioles resembling those of the unrelated Fabaceae. Quiinaceae. An obscure tropical family with three medium-sized tree species. Quiina has two species, one quite common, with opposite leaves and four stipules at each pair. Lac­ unaria has leaves in whorls of three. Rosaceae. The rose family is a large and popular family of the temperate zone plus tropical mountains. There are three tree species in Prunus, one in Photinia, and a marginal treelet in Hesperomeles. Photinia has whorled leaves; others are alternate. Prunus has untoothed leaves; others are toothed. Sabiaceae. A fairly important family we elected to omit. Meliosma has 12 mediumsized tree species with simple, alternate, toothed, clustered leaves, mostly in the mountains but also in lowland wet forests. Saxifragaceae. There are no longer trees in

477

Panama in this family. See Escalloniaceae and Phyllonomaceae. Schoepfiaceae. Two species of Schoepfia are small trees with simple, alternate leaves of montane forest. They used to be classified in the Olacaceae. Solanaceae. The nightshade family, including tomato and potato, is by far the biggest family we excluded. We count 60 species in 11 genera as trees, including 34 Solanum (the potato genus). Some are medium-sized trees, but most are small, and they grade into weedy shrubs. Stemonuraceae. Discophora guianensis is a fairly common, lowland, wet-forest tree species that we encounter often in the Canal Area and is widely known in Costa Rica. It has simple, alternate leaves, rather like those of Lauraceae. It was formerly classified in the Icacinaceae, which is covered in the text. Styracaceae. Four species of medium-sized trees in the mostly temperate snowbell genus, Styrax, are found in montane forest. They have simple, alternate leaves. Symplocaceae. The genus Symplocos includes seven tree species of montane forest (one reaches lowland wet forest). They have toothed, alternate, pubescent, two-ranked leaves. Theaceae. The tea family of Panama is now pared down to two species of Gordonia of montane and lowland wet forests. They have simple, alternate, toothed, bunched leaves and big, white, showy flowers. Other species formerly in the family have been moved to Ternstroemiaceae, which is covered in the text. Theophrastaceae. This family includes two genera, both with small, orange flowers and simple, alternate leaves. Jacquinia is a dryforest tree with spines on leaf tips. Clavija has three treelet species of wet-forest understory; two have very long leaves, like those of juvenile Gustavia (Lecythidaceae, covered in the text), whereas the third has short leaves. Ticodendraceae. A species discovered in 1989 in Costa Rica was so distinct that a new genus (Ticodendron) and a new family were created for it. It is a large tree resembling Alnus (Betulaceae, covered in the text),

478

Appendix 3

known best in montane forests of Costa Rica, but now collected in w. Panama too. Turneraceae. One species each in the genera Erblichia and Turnera are treelets with simple, alternate, toothed leaves. The latter resembles Hybanthus (Violaceae, covered in the text). Winteraceae. A single species of Drimys is

common in the mountains. It has simple, alternate leaves that are smooth and white below. Zamiaceae. Cycads are familiar garden plants that look like palms but are not related. About six species of Zamia with thick woody stems, reaching 2–3 m in height, are found in montane and lowland wet areas.

Literature Cited Barthlott, W., Lauer, W., Placke, A. 1996. Global distribution of species diversity in vascular plants: towards a world map of phytodiversity. Erdkunde 50: 317–327. Condit, R., Aguilar, S., Hernández, A., Pérez, R., Lao, S., Angehr, G., Hubbell, S.P., Foster, R.B. 2004. Tropical forest dynamics across a rainfall gradient and the impact of an El Niño dry season. Journal of Tropi­ cal Ecology 20: 51–72. Condit, R., Aguilar, S., Hernández, A., Pyke, C.R., Lao, S. 2005. Spatial changes in tree composition of high diversity forests: how much is predictable? Pp. 271–294 in Tropical Rainforests: Past, Present and Future (E. Bermingham, C. Dick, and C. Moritz, eds.). University of Chicago Press, Chicago, Illinois, USA. Condit, R., Pitman, N., Leigh, E.G., Chave, J., Terborgh, J., Foster, R.B., Núñez, P.V., Aguilar, S.,Valencia, R.,Villa, G., Muller-Landau, H., Losos, E., Hubbell, S.P. 2002. Beta-diversity in tropical forest trees. Science 295: 666–669. Condit, R., Robinson, W.D., Ibáñez, R., Aguilar, S., Sanjur, A., Martínez, R., Stallard, R., García, T., Angehr, G., Petit, L., Wright, S.J., Robinson, T.R., Heckadon, S. 2001. Maintaining the Canal while conserving biodiversity around it: a challenge for economic development in Panama in the 21st century. Bio­ Science 51: 135–144. Correa, M., Galdames, C., de Stapf, M.S. 2004. Catálogo de las Plantas Vasculares de Panamá. Quebecor World, Bogotá, Colombia. Croat, T.R. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, California, USA. Elias, T.S. 1980. The Complete Trees of North America. Van Nostrum Reinhold, New York, New York, USA. Engelbrecht, B.M.J., Comita, L.S., Condit, R., Kursar, T.A., Tyree, M.T., Turner, B.L., Hubbell, S.P. 2007. Drought sensitivity shapes species distribution patterns in tropical forests. Nature 447: 80–83.

Gentry, A.H. 1996. A Field Guide to the Families and Genera of Woody Plants of Northwest South America. University of Chicago Press, Chicago, Illinois, USA. Harris, J.G., Harris, M.W. 1994. Plant Identification Terminology. Spring Lake Publishing, Spring Lake, Utah, USA. Holdridge, L.R. 1967. Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica. Holdridge, L.R, Poveda A., L.J. 1975. Árboles de Costa Rica, vol. I. Centro Científo Tropical, San Jose, Costa Rica. Johnson, O., More, D. 2004. Tree Guide. Harper Collins, London, UK. Niehaus, T.F., Ripper, C.L. 1976. Field Guide to Pacific States Wildflowers. Houghton Mifflin Co., New York, New York, USA. Peterson, R.T. 1934. A Field Guide to the Birds. Houghton Mifflin Co., Boston, Massachusetts, USA. Peterson, R.T., McKenny, M. 1968. A Field Guide to Wildflowers. Houghton Mifflin Co., Boston, Massachusetts, USA. Petrides, G. 1972. A Field Guide to Trees and Shrubs. Houghton Mifflin Co., Boston, Massachusetts, USA. Ridgely, R. 1978. Field Guide to the Birds of Panama. Princeton University Press, Princeton, New Jersey, USA. Smith, N., Mori, S.A., Henderson, A., Stevenson, D.W., Heald, S.V. 2004. Flowering Plants of the Neotropics. Princeton University Press, Princeton, New Jersey, USA. Zamora V., N., Jiménez M., Q., Poveda A., L.J. 2000. Árboles de Costa Rica, vol. II. Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica. Zamora V., N., Jiménez M., Q., Poveda A., L.J. 2004. Árboles de Costa Rica, vol. III. Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica.

Index Latin names are italicized; common names are not. Where two names appear on a row, the second in parentheses is the name by which the species appears in the main text. If the first name is not italicized, it is a common name for the given species. If it is italicized, the first name is an out-dated and now invalid Latin name. The invalid names do not appear anywhere else in the book, but may be useful to readers who have studied Central American trees in the past and learned species by old names. Common names may appear more than once, in cases where the same name is applied to more than one species. Abarema barbouriana, 186 acacia africana (Peltophorum pterocarpum), 182 Acacia melanoceras, 186, 187 Acalypha diversifolia, 160, 161 aceituno (Simarouba amara), 448 achiote (Bixa orellana), 94 achiote (Vismia baccifera), 242 achiote (Vismia macrophylla), 244 achiote tigre (Vismia baccifera), 242 achiotillo (Alchornea latifolia), 164 Acosmium panamense, 214, 215 Adelia triloba, 162, 163 Aegiphila anomala, 248, 249 Aegiphila glandulifera (Aegiphila panamensis), 250 Aegiphila magnifica (Aegiphila panamensis), 250 Aegiphila odontophylla, 250, 251 Aegiphila panamensis, 250, 251 Aegiphila pendula (Aegiphila panamensis), 250 African tulip tree (Spathodea campanulata), 90 aguacate (Persea americana), 260 aguacatillo (Beilschmiedia pendula), 252 Aiouea lundelliana (Ocotea insularis), 260 ajicillo (Heisteria acuminata), 156 ajicillo (Heisteria concinna), 156 ajo (Cassipourea elliptica), 378 alazano (Calycophyllum candidissimum), 384 Albizia adinocephala, 188 Albizia guachapele (Pseudosamanea guachapele), 210 Albizia longepedata (Pseudosamanea guachapele), 210 alcanfor (Protium costaricense), 106 alcarreto (Aspidosperma spruceanum), 64 alcarreto (Erythroxylum citrifolium), 158

alcarreto (Erythroxylum macrophyllum), 158 alcarreto (Erythroxylum panamense), 160 Alchornea costaricensis, 162, 163 Alchornea latifolia, 164, 165 alcornoque (Ormosia coccinea), 230 alcornoque (Ormosia macrocalyx), 230 alfaje (Trichilia pleeana), 320 alfaje (Trichilia tuberculata), 320 alfajía (Trichilia pleeana), 320 alfajía (Trichilia tuberculata), 320 alfajía colorado (Trichilia tuberculata), 320 algarrobillo (Crudia acuminata), 180 algarrobillo (Lennea viridiflora), 224 algarrobillo (Prosopis juliflora), 210 algarrobo (Hymenaea courbaril), 180 algodoncillo (Croton draco), 166 Alibertia edulis, 380, 381 aliso (Alnus acuminata), 86 Allophylus psilospermus, 426, 427 almácigo (Bursera simaruba), 106 almendro (Dipteryx oleifera), 220 almendro de montaña (Dipteryx oleifera), 220 almendro de río (Andira inermis), 216 Alnus acuminata, 86, 87 Alnus ferruginea (Alnus acuminata), 86 Alseis blackiana, 382, 383 Amaioua corymbosa, 382, 383 Amanoa guianensis, 362, 363 amapola (Wercklea cocleana), 292 amapola (Wercklea woodsonii), 294 amargo-amargo (Vatairea erythrocarpa), 236 amarillo (Buchenavia tetraphylla), 144 amarillo (Bucida buceras), 146 amarillo (Lafoensia punicifolia), 268 amarillo (Maclura tinctoria), 326 amarillo (Terminalia amazonia), 148 amarillo carabazuelo (Terminalia amazonia), 148

amarillo de pepita (Buchenavia tetraphylla), 144 amarillo pepita (Lafoensia punicifolia), 268 American oil palm (Elaeis oleifera), 82 Amphitecna latifolia, 88, 89 Anacardium excelsum, 30, 31, 32 Anacardium occidentale, 32, 33 Anaxagorea panamensis, 38, 39 Andira inermis, 216, 217 animé (Tetragastris panamensis), 110 Annona glabra, 38, 39 Annona hayesii, 40, 41 Annona pittieri, 40, 41 Annona purpurea, 42, 43 Annona spraguei, 42, 43 annonillo (Rollinia mucosa), 54 annonillo (Rollinia pittieri), 56 anon de agua (Annona glabra), 38 anon de montaña (Annona pittieri), 40 anon de pantano (Annona glabra), 38 anon de puerco (Annona glabra), 38 Antirhea trichantha (Pittoniotis trichantha), 400 Apeiba aspera (Apeiba membranacea), 286 Apeiba membranacea, 286, 287 Apeiba tibourbou, 286, 287 Aphelandra sinclairiana, 26, 27 Arachnothryx buddleioides, 384, 385 árbol carne (Roupala montana), 372 árbol caspa (Zuelania guidonia), 424 árbol del vela (Parmentiera cereifera), 90 árbol sarigua (Parkinsonia aculeata), 182 arbol soga (Platypodium elegans), 232 arcabú (Zanthoxylum acuminatum), 412 arcabú (Zanthoxylum ekmanii), 412 arcabú (Zanthoxylum panamense), 412

482

Index

arcabú (Zanthoxylum setulosum), 414 Ardisia acutata (Ardisia standleyana), 346 Ardisia bartlettii, 344, 345 Ardisia fendleri (Ardisia standleyana), 346 Ardisia polyantha (Ardisia standleyana), 346 Ardisia polydactyla (Ardisia standleyana), 346 Ardisia revoluta, 346, 347 Ardisia standleyana, 346, 347 aromo (Prosopis juliflora), 210 Aspidosperma cruentum (Aspidosperma spruceanum), 64 Aspidosperma megalocarpon, 62 Aspidosperma spruceanum, 64, 65 Astrocaryum standleyanum, 78, 79 Astronium graveolens, 32, 33 Attalea butyracea, 80, 81 Avicennia bicolor, 26, 27 Avicennia germinans, 28, 29 Avicennia tonduzii (Avicennia germinans), 28 avocado (Persea americana), 260 azote (Hampea appendiculata), 292 azulejo (Coutarea hexandra), 388 azulejo (Exostema mexicanum), 388 baco (Magnolia sororum), 268 Bactris augustinea (Bactris major), 80 Bactris balanoidea (Bactris major), 80 Bactris major, 80 Bactris superior (Bactris major), 80 bagre (Adelia triloba), 162 bala de cañón (Couroupita guianensis), 262 balata (Manilkara bidentata), 438 balo (Gliricidia sepium), 224 balsa (Ochroma pyramidale), 276 bálsamo (Myroxylon balsamum), 228 bálsamo de tolú (Myroxylon balsamum), 228 balso (Ochroma pyramidale), 276 Banara guianensis, 414, 415 barbaso (Senna dariensis), 184 barillo (Symphonia globulifera), 140 barrigón (Pseudobombax septenatum), 280 Beilschmiedia pendula, 252, 253 Bellucia axinanthera (Bellucia pentamera), 296 Bellucia costaricensis (Bellucia pentamera), 296 Bellucia pentamera, 296, 297 benjamín (Faramea luteovirens), 390

benjamín (Faramea occidentalis), 390 berbá (Brosimum alicastrum), 322 berbá (Brosimum guianense), 322 Bixa orellana, 94, 95 Bixa urucurana (Bixa orellana), 94 biyuyo (Cordia dentata), 100 black mangrove (Avicennia bicolor), 26 black mangrove (Avicennia germinans), 28 black palm (Astrocaryum standleyanum), 78 boca de vieja (Posoqueria latifolia), 402 Bocconia frutescens, 362, 363 bogamani (Virola sebifera), 342 bogamani (Virola surinamensis), 344 Bombacopsis quinata (Pachira quinata), 278 Bombacopsis sessilis (Pachira sessilis), 278 Bonafousia panamensis (Tabernaemontana panamensis), 72 Bonafousia undulata (Tabernaemontana undulata), 74 borojó (Posoqueria latifolia), 402 borojocito (Rudgea pittieri), 408 botarrama (Vochysia ferruginea), 462 Bourreria costaricensis, 96, 97 Bourreria panamensis (Bourreria costaricensis), 96 breadnut (Brosimum alicastrum), 322 Brosimum alicastrum, 322, 323 Brosimum bernadetteae (Brosimum alicastrum), 322 Brosimum guianense, 322, 323 Brosimum latifolium (Brosimum alicastrum), 322 Brosimum utile, 324, 325 Brownea macrophylla, 176, 177 Buchenavia capitata (Buchenavia tetraphylla),144 Buchenavia tetraphylla, 144, 145 buchirago (Ladenbergia macrocarpa), 396 Bucida buceras, 146, 147 Buddleja americana, 446, 447 bull thorn (Acacia melanoceras), 186 Bunchosia cornifolia (Bunchosia nitida), 270 Bunchosia nitida, 270, 271 burrilico (Trichospermum galeottii), 290 Bursera simaruba, 106, 107

button mangrove (Conocarpus erectus), 146 Byrsonima crassifolia, 270, 271 Byrsonima spicata, 272, 273 cabeza de víbora (Faramea papirifolia), 392 cabimo (Copaifera aromatica), 178 cabresto (Ormosia coccinea), 230 cabresto (Ormosia macrocalyx), 230 cacao (Theobroma cacao), 284 cacao de monte (Herrania nycterodendron), 282 cacao de monte (Herrania purpurea), 284 cachito (Acacia melanoceras), 186 cachos de venado (Xylosma chlorantha), 424 cacique (Brosimum alicastrum), 322 cacique (Brosimum guianense), 322 cacique (Diphysa americana), 218 Caesalpinia coriaria, 176, 177 cafecillo (Psychotria grandis), 404 cafecillo (Psychotria horizontalis), 404 cafecillo (Psychotria luxurians), 406 cafecillo (Psychotria marginata), 406 cagajón (Zuelania guidonia), 424 caimito (Chrysophyllum cainito), 436 caimito de mono (Chrysophyllum argenteum), 434 caimito de montaña (Ecclinusa lanceolata), 438 calaba (Calophyllum longifolium), 132 calaba (Calophyllum nubicola), 134 calabacito (Amphitecna latifolia), 88 calabacito (Lafoensia punicifolia), 268 calabazo (Crescentia cujete), 88 Calatola costaricensis, 246, 247 Calatola microcarpa (Calatola costaricensis), 246 Calliandra magdalenae, 188 Calophyllum longifolium, 132, 133 Calophyllum nubicola, 134, 135 Calycolpus warszewiczianus, 350, 351 Calycophyllum candidissimum, 384, 385 camaron (Hirtella triandra), 124 camaron (Hirtella tubiflora), 126 camaroncillo (Hirtella americana), 124 camaroncillo (Hirtella triandra), 124 camaroncillo (Hirtella tubiflora), 126 Campnosperma panamense, 34 caña brava (Bactris major), 80 caña fistula (Cassia fistula), 178 caña fístula (Cassia moschata), 178

Index canaleto (Macrocnemum roseum), 398 canaleto (Platypodium elegans), 232 canalú (Bourreria costaricensis), 96 canalua (Platypodium elegans), 232 candelabro (Cespedesia spathulata), 360 candelillo (Cupania rufescens), 428 candelo (Pittoniotis trichantha), 400 canelito (Ardisia bartlettii), 344 canelito (Ardisia revoluta), 346 canelito (Ardisia standleyana), 346 canelito (Hamelia patens), 396 canelito (Neea amplifolia), 358 canelito (Stylogyne turbacensis), 350 canelo (Annona hayesii), 40 canelo (Ardisia bartlettii), 344 canelo (Ardisia revoluta), 346 canelo (Ardisia standleyana), 346 canelo (Neea amplifolia), 358 canillo (Clidemia octona), 298 canillo (Conostegia bracteata), 298 canillo (Conostegia cinnamomea), 298 canillo (Conostegia rufescens), 300 canillo (Conostegia speciosa), 300 canillo (Henriettea succosa), 302 canillo (Miconia affinis), 302 canillo (Miconia centronioides), 304 canillo (Miconia elata), 304 canillo (Miconia hondurensis), 306 canillo (Miconia nervosa), 308 canillo (Miconia trinervia), 308 cannon-ball tree (Couroupita guianensis), 262 caoba (Swietenia macrophylla), 316 caobilla (Tapirira guianensis), 36 Capparis baduca (Capparis frondosa), 114 Capparis crotonantha (Capparis pittieri), 116 Capparis cynophallophora, 114, 115 Capparis frondosa, 114, 115 Capparis indica, 116, 117 Capparis isthmensis (Capparis cynophallophora), 114 Capparis pittieri, 116, 117 capulín (Muntingia calabura), 334 capulín (Trema micrantha), 112 capulín (Trichospermum galeottii), 290 caracucha (Plumeria rubra), 68 caraño (Trattinnickia aspera), 110 caraño (Zuelania guidonia), 424 carao (Cassia moschata), 178 Carapa guianensis, 310, 311

Carapa nicaraguensis (Carapa guianensis), 310 Carapa slateri (Carapa guianensis), 310 carapelo (Couratari guianensis), 262 carbonero (Colubrina glandulosa), 374 carbonero (Lindackeria laurina), 28 carcuera (Platypodium elegans), 232 Carica papaya, 118 carne asada (Roupala montana), 372 carne de venado (Capparis cynophallophora), 114 carne de venado (Capparis indica), 116 Caryodaphnopsis burgeri, 254, 255 casaco (Sloanea terniflora), 154 Casearia arborea, 416, 417 Casearia arguta, 416, 417 Casearia commersoniana, 418, 419 Casearia sylvestris, 418, 419 cashew (Anacardium occidentale), 32 casia amarilla (Cassia moschata), 178 casiquillo (Mabea occidentalis), 170 Cassia fistula, 178 Cassia fruticosa (Senna dariensis), 184 Cassia moschata, 178, 179 Cassia reticulata (Senna reticulata), 184 Cassipourea elliptica, 378, 379 Castilla elastica, 324 cativo (Prioria copaifera), 182 cauchillo (Sorocea affinis), 332 caucho (Castilla elastica), 324 caucho (Hevea brasiliensis), 166 Cavanillesia platanifolia, 274, 275 Cecropia arachnoidea (Cecropia peltata), 458 Cecropia eximia (Cecropia insignis), 454 Cecropia garciae, 454 Cecropia insignis, 454, 455 Cecropia longipes, 456, 457 Cecropia maxonii (Cecropia obtusifolia), 456 Cecropia obtusifolia, 456, 457 Cecropia panamensis (Cecropia obtusifolia), 456 Cecropia peltata, 458, 459 Cedrela odorata, 312, 313 cedrillo (Turpinia occidentalis), 450 cedrillo de montaña (Turpinia occidentalis), 450 cedro (Cedrela odorata), 312 cedro amargo (Cedrela odorata), 312

483

cedro bateo (Carapa guianensis), 310 cedro espino (Pachira quinata), 278 cedro macho (Guarea grandifolia), 312 cedro macho (Guarea guidonia), 314 Ceiba pentandra, 274, 275 ceibo (Ceiba pentandra), 274 ceibo (Hura crepitans),168 ceibo (Pachira sessilis), 278 Celtis schippii, 112, 113 cenízaro (Samanea saman), 212 cenizo (Licania affinis), 126 cenizo (Ulmus mexicana), 454 Cephaelis barcellana (Psychotria poeppigiana), 408 Cephaelis elata (Psychotria elata), 402 Cephaelis tomentosa (Psychotria poeppigiana), 408 Cephaelis vultusmimi (Psychotria poeppigiana), 408 cerezo de monte (Bunchosia nitida), 270 cerillo (Perebea xanthochyma), 330 cerillo (Symphonia globulifera), 140 cero (Symphonia globulifera), 140 Cespedesia macrophylla (Cespedesia spathulata), 360 Cespedesia spathulata, 360, 361 Cespedezia macrophylla (Cespedesia spathulata), 360 Chamaedorea tepejilote, 80, 81 Chamaedorea wedlandiana (Chamaedorea tepejilote), 80 Chamaedorea wendlandiana (Chamaedorea tepejilote), 80 chaparrón (Garcinia intermedia), 136 chaparrón (Garcinia madruno), 136 chaperno (Lonchocarpus heptaphyllus), 224 chaperno (Lonchocarpus sericeus), 226 chaperno (Lonchocarpus velutinus), 226 chicarrón (Hirtella triandra), 124 chiricano (Vantanea depleta), 242 chirimoya (Annona spraguei), 42 chirimoya (Rollinia mucosa), 54 Chlorophora tinctoria (Maclura tinctoria), 326 chocolate (Theobroma cacao), 284 chorola (Heisteria acuminata), 156 chorola (Heisteria concinna), 156 Chrysophyllum argenteum 434, 435 Chrysophyllum cainito, 436, 437 Chrysophyllum colombianum, 436, 437

484

Index

Chrysophyllum panamense (Chrysophyllum argenteum), 434 chuchupate (Guarea grandifolia), 312 chuchupate (Guarea guidonia), 314 chumico (Curatella americana), 152 chumico sabanero (Curatella americana), 152 chunga (Astrocaryum standleyanum), 78 chutra (Protium costaricense), 106 chutra (Protium panamense), 108 chutra (Protium tenuifolium), 108 chutra (Tetragastris panamensis), 110 cierito (Mouriri myrtilloides), 310 cigarrillo (Schizolobium parahyba), 184 Cinnamomum cinnamomifolium (Cinnamomum triplinerve), 254 Cinnamomum triplinerve,254, 255 Citharexylum caudatum, 460, 461 Citrus sinensis,410 clavito (Margaritaria nobilis), 366 clavito (Pera arborea), 172 Clethra lanata, 132, 133 Clidemia octona, 298, 299 Clitoria glaberrima, 216, 217 Clusia pratensis, 134, 135 Coccoloba manzinellensis, 368, 369 Coccoloba uvifera, 370, 371 Cochlospermum vitifolium, 144, 145 coco (Couroupita guianensis), 262 coco (Lecythis ampla), 266 coco de agua (Pachira aquatica), 276 cocobolo (Dalbergia retusa), 218 Cojoba membranaceum (Cojoba rufescens), 188 Cojoba rufescens, 188, 189 Cojoba tubulifera (Cojoba rufescens), 188 Cojoba tubuliferum (Cojoba rufescens), 188 Cojoba whitefoordiae (Cojoba rufescens), 188 Colubrina glandulosa, 374, 375 Colubrina heteroneura, 376, 377 Compsoneura capitellata, 336, 337 Compsoneura sprucei,338, 339 condón de mono (Couratari guianensis), 262 conejito colorado (Ormosia macrocalyx), 230 conejito colorado (Trichilia pallida), 318 conejo (Hirtella triandra), 124 conejo (Laetia thamnia), 422

conejo colorado (Laetia thamnia), 422 conejo colorado (Trichilia hirta), 316 conejo colorado (Trichilia martiana), 318 conga (Welfia regia), 84 congolo (Couratari guianensis), 262 Conocarpus erectus, 146, 147 Conostegia bracteata, 298 Conostegia cinnamomea, 298, 299 Conostegia micromeris (Conostegia cinnamomea), 298 Conostegia puberula (Conostegia rufescens), 300 Conostegia rufescens, 300 Conostegia speciosa, 300, 301 Conostegia xalapensis,300, 301 Copaifera aromatica, 178, 179 copal (Protium costaricense), 106 copal (Protium panamense), 108 copal (Protium tenuifolium), 108 copé (Clusia pratensis), 134 copidijo (Virola sebifera), 342 coquillo (Jatropha curcas), 170 coquito (Couratari guianensis), 262 coquito (Eschweilera pittieri), 264 coquito (Eschweilera sessilis), 264 coralillo (Cojoba rufescens), 188 coralillo (Ormosia amazonica), 228 coralillo (Ormosia coccinea), 230 coralillo (Ormosia macrocalyx), 230 Cordia alliodora, 96, 97 Cordia bicolor, 98, 99 Cordia cymosa, 98, 99 Cordia dentata, 100, 101 Cordia dwyeri, 100, 101 Cordia eriostigma, 102, 103 Cordia lasiocalyx, 102, 103 Cordia panamensis, 104, 105 Cordia porcata, 104, 105 corneto (Iriartea deltoidea), 82 Cornus disciflora, 150, 151 corocillo (Licania hypoleuca), 128 corocito (Humiriastrum diguense), 240 corocito (Maranthes panamensis), 130 corocito (Sacoglottis trichogyna), 240 corocito (Vantanea depleta), 242 coronillo (Bellucia pentamera), 296 corotú (Enterolobium cyclocarpum), 188 corotú de montaña (Enterolobium schomburgkii), 190 corozo (Bactris major), 80 corozo (Elaeis oleifera), 82

corozo (Humiriastrum diguense), 240 corozo (Maranthes panamensis), 130 corozo (Sacoglottis trichogyna), 240 corozo colorado (Elaeis oleifera), 82 Corozo oleifera (Elaeis oleifera), 82 corta lengua (Banara guianensis), 414 corta lengua (Casearia arborea), 416 corta lengua (Casearia commersoniana), 418 corta lengua (Casearia sylvestris), 418 corta lengua (Hasseltia floribunda), 420 corta lengua (Ryania speciosa), 422 cortezo (Apeiba membranacea), 286 cortezo (Apeiba tibourbou), 286 Cosmibuena macrocarpa 386, 387 Cosmibuena paludicola (Cosmibuena macrocarpa), 386 costilla (Platypodium elegans), 232 Couepia panamensis (Maranthes panamensis), 130 Couma macrocarpa, 64 Coumarouna oleifera (Dipteryx oleifera), 220 Couratari guianensis, 262, 263 Couratari panamensis (Couratari guianensis), 262 Couroupita guianensis, 262, 263 Couroupita idolica (Couroupita guianensis), 262 Coussarea curvigemmia, 386, 387 Coutarea hexandra, 388, 389 Crateva tapia, 118, 119 Cremastosperma sp. (Mosannona garwoodii), 52 Crescentia cujete, 88, 89 cresta de gallo (Pogonopus exsertus), 400 cresta de gallo (Warszewiczia coccinea), 410 cricamola (Pterocarpus officinalis), 234 cricamola (Pterocarpus rohrii), 234 criollo (Minquartia guianensis), 158 criollo negro (Minquartia guianensis), 158 Crossopetalum eucymosum (Crossopetalum parviflorum), 120 Crossopetalum parviflorum, 120, 121 Croton billbergianus, 164, 165 Croton draco, 166, 167 Croton panamensis (Croton draco), 166 cruceta (Quassia amara), 448 Crudia acuminata, 180, 181

Index Cryosophila albida (Cryosophila warscewiczii), 80 Cryosophila warscewiczii, 80 cuajado (Minquartia guianensis), 158 cuajado (Vitex cooperi), 252 cuajado negro (Minquartia guianensis), 158 cuatro estomagos (Tetragastris panamensis), 110 cucaracho (Astronium graveolens), 32 cucharo (Schizolobium parahyba), 184 cuchillito (Brownea macrophylla), 176 cucua (Poulsenia armata), 330 cuernito (Acacia melanoceras), 186 cuipo (Cavanillesia platanifolia), 274 Cupania cinerea, 426, 427 Cupania latifolia, 428, 429 Cupania rufescens, 428, 429 Cupania scrobiculata, 430 Cupania seemannii, 430, 431 Cupania sylvatica (Cupania seemannii), 430 Curatella americana, 152, 153 cutarro (Swartzia panamensis), 234 Cymbopetalum lanugipetalum, 44, 45 Cynodendron panamense (Chrysophyllum argenteum), 434 Cynometra bauhiniifolia, 180 Cyrilla racemiflora, 152, 153 Dalbergia retusa, 218, 219 dandruff tree (Zuelania guidonia), 424 Daphnopsis americana, 452, 453 Daphnopsis seiberti (Daphnopsis americana), 452 dardo (Ruprechtia costata), 370 Dendropanax acreanum (Dendropanax arboreus), 76 Dendropanax arboreus, 76, 77 Dendropanax stenodontus (Dendropanax arboreus), 76 Dendrosicus latifolius (Amphitecna latifolia), 88 Desmopsis panamensis, 44, 45 Dialyanthera acuminata (Otoba acuminata), 338 Dialyanthera otoba (Otoba novogranatensis), 340 Didymopanax morototoni (Schefflera morototoni), 76 Diospyros artanthifolia, 154, 155 Diphysa americana, 218, 219

Diphysa robinioides (Diphysa americana), 218 Diplotropis purpurea, 220, 221 Dipteryx oleifera, 220, 221 Dipteryx panamensis (Dipteryx oleifera), 220 dormilón (Enterolobium schomburgkii), 190 dormilón (Parkia nitida), 208 dos caras (Arachnothryx buddleioides), 384 dos caras (Conostegia xalapensis), 300 dos caras (Miconia argentea), 304 dos caras (Miconia centronioides), 304 dos caras (Miconia elata), 304 dos caras (Miconia impetiolaris), 306 Drypetes standleyi, 374, 375 Duguetia confusa, 46, 47 ear tree (Enterolobium cyclocarpum), 188 Ecclinusa lanceolata, 438, 439 Elaeis oleifera, 82, 83 elm (Ulmus mexicana), 454 encino (Quercus insignis), 238 Enterolobium cyclocarpum,188, 189 Enterolobium schomburgkii, 190, 191 Erythrina costaricensis, 222, 223 Erythrina fusca, 222, 223 Erythroxylum citrifolium, 158, 159 Erythroxylum lucidum (Erythroxylum macrophyllum), 158 Erythroxylum macrophyllum, 158, 159 Erythroxylum multiflorum (Erythroxylum macrophyllum), 158 Erythroxylum panamense, 160, 161 Erythroxylum skutchii (Erythroxylum macrophyllum), 158 Eschweilera pittieri, 264 Eschweilera reversa (Eschweilera pittieri), 264 Eschweilera sessilis, 264, 265 Eschweilera verruculosa (Eschweilera pittieri), 264 espavé (Anacardium excelsum), 30 espino amarillo (Adelia triloba), 162 espino amarillo (Guettarda foliacea), 394 espino carbón (Pithecellobium unguis­cati), 208 espino del diablo (Colubrina heteroneura), 376 espino rico (Sideroxylon celastrinum), 446

485

esquitilla (Allophylus psilospermus), 426 esquitillo (Allophylus psilospermus), 426 Eugenia coloradoensis, 352, 353 Eugenia nesiotica, 352, 353 Eugenia oerstediana, 352 Eugenia principium, 354, 355 Exostema mexicanum, 388, 389 faldita de puta (Pouteria reticulata), 442 faldita de puta (Pouteria stipitata), 444 Faramea caput­anguis (Faramea papirifolia), 392 Faramea luteovirens, 390, 391 Faramea occidentalis, 390, 391 Faramea papirifolia, 392, 393 felí (Pera arborea), 172 Ficus glabrata (Ficus insipida), 324 Ficus insipida, 324, 325 Ficus tonduzii, 326, 327 fig (Ficus insipida), 324 fig (Ficus tonduzii), 326 flor azul (Vitex cooperi), 252 flor de mayo (Vochysia ferruginea), 462 flores de lluvia (Hybanthus prunifolius), 460 fosforito (Trichilia pleeana), 320 fosforito (Trichilia tuberculata), 320 frangipani (Plumeria rubra), 68 frijolillo (Acosmium panamense), 214 frijolillo (Lonchocarpus minimiflorus), 226 frijolillo (Pseudosamanea guachapele), 210 frijolito de la suerte (Ormosia amazonica), 228 frijolito de la suerte (Ormosia coccinea), 230 frio (Colubrina glandulosa), 374 fruta de mono (Aegiphila panamensis), 250 fruta de mono (Ladenbergia macrocarpa), 396 fruta de mono (Posoqueria latifolia), 402 fruta dorada (Otoba novogranatensis), 340 fruta dorada (Virola koschnyi), 340 fruta dorada (Virola surinamensis), 344 fruto de mono (Rudgea pittieri), 408

486

Index

gallinazo (Schizolobium parahyba), 184 gallito (Calliandra magdalenae), 188 gallito (Erythrina fusca), 222 gallote (Sorocea affinis), 332 gallote (Trophis racemosa), 334 Garcinia acuminata (Garcinia madruno), 136 Garcinia intermedia, 136, 137 Garcinia madruno, 136, 137 garrapato (Hirtella americana), 124 garrapato (Hirtella triandra), 124 garrapato (Licania hypoleuca), 128 garrotillo (Capparis frondosa), 114 garrotillo (Faramea luteovirens), 390 garrotillo (Faramea occidentalis), 390 garrotillo (Lozania pittieri), 248 Genipa americana, 392, 393 Geoffroea inermis (Andira inermis), 216 Geonoma cuneata, 82 Geonoma decurrens (Geonoma cuneata), 82 Geonoma gracilis (Geonoma cuneata), 82 Geonoma obovata (Geonoma cuneata), 82 Geonoma procumbens (Geonoma cuneata), 82 Gliricidia sepium, 224, 225 gorgojero (Cupania cinerea), 426 gorgojero (Cupania latifolia), 428 gorgojero (Cupania rufescens), 428 gorgojero (Cupania seemannii), 430 gorgojero (Matayba apetala), 430 gorgojo (Cupania latifolia), 428 gorgojo (Cupania rufescens), 428 gorgojo (Cupania seemannii), 430 gorgojo (Matayba apetala), 430 gorgojo blanco (Cupania cinerea), 426 Graphardisia bartlettii (Ardisia bartlettii), 344 Graphardisia lewisii (Ardisia bartlettii), 344 Graphardisia romeroi (Ardisia bartlettii), 344 Graphardisia sapoana (Ardisia bartlettii), 344 Graphardisia tuirana (Ardisia bartlettii), 344 guaba (all Inga species), 190-206 guabanday (Jacaranda copaia), 90 guábilo (Enterolobium schomburgkii), 190

guábilo (Pseudosamanea guachapele), 210 guabito (all Inga species), 190-206 guabito (Lonchocarpus sericeus), 226 guabito amargo (Quassia amara), 448 guabito cansa boca (Zygia latifolia), 212 guabito cansa boca (Zygia longifolia), 214 guabito de río (Zygia latifolia), 212 guabito de río (Zygia longifolia), 214 guabo (all Inga species), 190-206 guachapalí (Pseudosamanea guachapele), 210 guachapalí (Samanea saman), 212 guácimo (Guazuma ulmifolia), 282 guácimo blanco (Luehea speciosa), 290 guácimo borcico (Luehea speciosa), 290 guácimo colorado (Luehea seemannii), 288 guácimo molenillo (Luehea seemannii), 288 guácimo molenillo (Luehea speciosa), 290 guácimo pacheco (Luehea seemannii), 288 guácimo tortugo (Luehea speciosa), 290 guaco (Crateva tapia), 118 guágara (Cryosophila warscewiczii), 80 guanábana de pantano (Annona glabra), 38 guanacaste (Enterolobium cyclocarpum), 188 guapinol (Hymenaea courbaril), 180 Guapira standleyana, 358, 359 Guarea grandifolia, 312, 313 Guarea guidonia, 314, 315 Guarea pterorhachis, 314, 315 guarumo (Cecropia insignis), 454 guarumo (Cecropia longipes), 456 guarumo (Cecropia obtusifolia), 456 guarumo (Cecropia peltata), 458 guarumo macho (Pourouma bicolor), 458 Guatteria aberrans, 46, 47 Guatteria amplifolia, 48, 49 Guatteria dumetorum, 48, 49 Guatteria jefensis, 50 Guatteria recurvisepala, 50, 51 Guatteria sessilicarpa, 50, 51 guava (Psidium guajava), 356 guayaba (Psidium guajava), 356

guayabillo (Calycolpus warszewiczianus), 350 guayabillo (Eugenia oerstediana), 352 guayabillo (Eugenia principium), 354 guayabillo (Mouriri myrtilloides), 310 guayabillo (Quararibea asterolepis), 280 guayabillo (Terminalia oblonga), 150 guayabita sabanera (Psidium guineense), 356 guayabito (Eugenia coloradoensis), 352 guayabito de montaña (Eugenia coloradoensis), 352 guayabito de monte (Alibertia edulis), 380 guayabito de monte (Calycolpus warszewiczianus), 350 guayabo de montaña (Terminalia oblonga), 150 guayabo de monte (Guettarda foliacea), 394 guayabo de sabana (Psidium guineense), 356 guayabo hormiguero (Triplaris cumingiana), 372 guayabo negro (Hamelia patens), 396 guayabón (Terminalia oblonga), 150 guayacan (Tabebuia guayacan), 92 guayacan (Tabebuia ochracea), 92 Guazuma ulmifolia, 282, 283 Guettarda chiriquiensis (Guettarda crispiflora), 394 Guettarda crispiflora, 394, 395 Guettarda foliacea, 394, 395 guindo (Ziziphus mauritiana), 378 gumbo limbo (Bursera simaruba), 106 Gustavia superba, 264, 265 Gyrocarpus americanus, 238 Hamelia patens, 396, 397 Hampea appendiculata, 292, 293 harino (Andira inermis), 216 harino (Cojoba rufescens), 188 harino (Enterolobium schomburgkii), 190 Hasseltia floribunda, 420, 421 Hasseltia gossypiosperma (Hasseltia floribunda), 420 Hasseltia panamensis (Hasseltia floribunda), 420 Hasseltia rigida (Hasseltia floribunda), 420 havillo (Hura crepitans), 168 Hedyosmum bonplandianum,120, 121

Index Hedyosmum calloso­serratum (Hedyosmum bonplandianum), 120 Hedyosmum costaricense, 122, 123 Hedyosmum mexicanum, 122, 123 Heisteria acuminata, 156, 157 Heisteria concinna, 156, 157 Heisteria cyanocarpa (Heisteria acuminata), 156 Heisteria longipes (Heisteria acuminata), 156 helicóptero (Gyrocarpus americanus), 238 Heliocarpus americanus, 288, 289 Heliocarpus popayanensis (Heliocarpus americanus), 288 Henriettea succosa, 302, 303 Hernandia didymantha, 238, 239 Herrania nycterodendron, 282, 283 Herrania purpurea, 284, 285 herrero (Mimosa tenuiflora), 206 Heterotrichum octonum (Clidemia octona), 298 Hevea brasiliensis, 166, 167 Hibiscus cocleanus (Wercklea cocleana), 292 Hibiscus woodsonii (Wercklea woodsonii), 294 Hieronyma alchorneoides, 364, 365 Hieronyma laxiflora (Hieronyma alchorneoides), 364 Hieronyma oblonga, 364, 365 higuerón (Ficus insipida), 324 higuerón (Ficus tonduzii), 326 hinojo (Piper reticulatum), 366 Hippomane mancinella, 168, 169 Hirtella americana, 124, 125 Hirtella triandra, 124, 125 Hirtella tubiflora, 126, 127 hombre grande (Quassia amara), 448 Hortia colombiana, 410, 411 hot lips (Psychotria poeppigiana), 408 huesito (Cassipourea elliptica), 378 huesito (Coussarea curvigemmia), 386 huesito (Faramea luteovirens), 390 huesito (Faramea occidentalis), 390 huesito (Lacistema aggregatum), 246 huesito (Lozania pittieri), 248 hueso (Coccoloba manzinellensis), 368 huevos de gato (Stemmadenia grandiflora), 70 huevos de gato (Stemmadenia obovata), 70 huevos de gato (Tabernaemontana arborea), 72

huevos de gato (Tabernaemontana panamensis), 72 huevos de gato (Tabernaemontana undulata), 74 huevos de gato (Thevetia ahouai), 74 hule (Castilla elastica), 324 Humiriastrum diguense, 240, 241 Hura crepitans, 168, 169 Hybanthus prunifolius, 460, 461 Hymenaea courbaril, 180, 181 Icacorea santafeana (Ardisia standleyana), 346 Icacorea standleyana (Ardisia standleyana), 346 ictericia (Senna dariensis), 184 indio (Schizolobium parahyba), 184 indio desnudo (Bursera simaruba), 106 Inga aestuariorum (Inga multijuga), 196 Inga affinis (Inga vera), 206 Inga alopetiolata (Inga sapindoides), 202 Inga bella, 190, 191 Inga chocoensis, 192, 193 Inga chriquensis (Inga edulis), 194 Inga chriquensis (Inga oerstediana), 198 Inga cocleensis, 192, 193 Inga confusa (Inga laurina), 194 Inga confusa (Inga ruiziana), 200 Inga coprocarpa (Inga chocoensis), 192 Inga edulis, 194 Inga edulis (Inga oerstediana), 198 Inga fagifolia (Inga laurina), 194 Inga fagifolia (Inga ruiziana), 200 Inga goldmanii,194, 195 Inga laurina,194, 195 Inga laurina (Inga ruiziana), 200 Inga lindeniana (Inga sapindoides), 202 Inga marginata, 196, 197 Inga microstachya (Inga pezizifera), 200 Inga minutula (Inga edulis), 194 Inga minutula (Inga oerstediana), 198 Inga mucuna, 196, 197 Inga multijuga, 196 Inga myriantha (Inga umbellifera), 204 Inga nobilis, 198, 199 Inga oerstediana, 198, 199 Inga oerstediana (Inga edulis), 194 Inga panamensis (Inga sapindoides), 202

487

Inga pavoniana (Inga sapindoides), 202 Inga pezizifera, 200, 201 Inga punctata, 200, 201 Inga quaternata (Inga nobilis), 198 Inga roussoviana (Inga nobilis), 198 Inga ruiziana, 200 Inga ruiziana (Inga laurina), 194 Inga sapindoides, 202, 203 Inga semialata (Inga marginata), 196 Inga spectabilis, 202, 203 Inga spuria (Inga vera), 206 Inga thibaudiana, 204, 205 Inga tysonii (Inga marginata), 196 Inga umbellifera, 204, 205 Inga urabensis (Inga venusta), 206 Inga venusta, 206, 207 Inga vera, 206 Inga williamsii (Inga nobilis), 198 Iriartea corneto (Iriartea deltoidea), 82 Iriartea deltoidea, 82, 83 Iriartea gigantea (Iriartea deltoidea), 82 jaboncillo (Sapindus saponaria), 432 jacaranda (Jacaranda copaia), 90 Jacaranda copaia, 90, 91 Jacaratia spinosa, 118, 119 jagua (Genipa americana), 392 jagua macho (Randia armata), 408 jaguey (Calatola costaricensis), 246 janeiro (Ormosia macrocalyx), 230 jarino (Enterolobium schomburgkii), 190 Jatropha curcas, 170, 171 jaúl (Alnus acuminata), 86 jazmín (Faramea luteovirens), 390 jazmín (Faramea occidentalis), 390 jira (Socratea exorrhiza), 84 jobo (Spondias mombin), 34 jobo de lagarto (Sciadodendron excelsum), 78 jordancillo (Trema micrantha), 112 kapok (Ceiba pentandra), 274 kerosín (Protium costaricense), 106 kerosín (Tetragastris panamensis), 110 labios ardientes (Psychotria poeppigiana), 408 labios de puta (Psychotria poeppigiana), 408 Lacistema aggregatum, 246, 247 Lacmellea panamensis, 64, 65 Lacmellea speciosa, 66, 67 Ladenbergia macrocarpa, 396, 397

488

Index

Laetia procera, 420, 421 Laetia thamnia, 422, 423 Lafoensia punicifolia, 268, 269 lagarto negro (Lacmellea panamensis), 64 Laguncularia racemosa, 148, 149 largarto (Zanthoxylum acuminatum), 412 largarto (Zanthoxylum panamense), 412 laso (Matayba scrobiculata), 432 laso prieto (Matayba scrobiculata), 432 laurel (Cordia alliodora), 96 laurel negro (Cordia cymosa), 98 laurel negro (Cordia dwyeri), 100 laureño (Senna reticulata), 184 Laxoplumeria tessmannii, 66 leche de vaca (Lacmellea panamensis), 64 leche de vaca (Lacmellea speciosa), 66 lechosa (Malouetia isthmica), 66 lechosa (Rauvolfia littoralis), 68 lechosa (Sorocea affinis), 332 lechosa (Trophis racemosa), 334 Lecythis ampla, 266, 267 lengua de vaca (Cordia panamensis), 104 Lennea viridiflora, 224 Licania affinis, 126, 127 Licania arborea, 128, 129 Licania hypoleuca, 128, 129 Licania platypus, 130, 131 lija (Trophis caucana), 332 limón de montaña (Capparis pittieri), 116 limón de montaña (Hortia colombiana), 410 limoncillo (Cassipourea elliptica), 378 limoncillo (Crossopetalum parviflorum), 120 limoncillo (Hedyosmum bonplandianum), 120 limoncillo (Siparuna pauciflora), 450 limoncillo (Swartzia simplex), 236 Lindackeria laurina, 28, 29 llanto (Guapira standleyana), 358 lloró (Cornus disciflora), 150 lluvia de plata (Calycophyllum candidissimum), 384 Lonchocarpus heptaphyllus, 224, 225 Lonchocarpus latifolius (Lonchocarpus heptaphyllus), 224 Lonchocarpus macrophyllus (Lonchocarpus sericeus), 226

Lonchocarpus minimiflorus, 226, 227 Lonchocarpus pentaphyllus (Lonchocarpus heptaphyllus), 224 Lonchocarpus sericeus, 226, 227 Lonchocarpus velutinus, 226 Lozania pedicellata (Lozania pittieri), 248 Lozania pittieri, 248, 249 Luehea seemannii, 288, 289 Luehea speciosa, 290, 291 Mabea occidentalis, 170, 171 macano (Diphysa americana), 218 machetito (Erythrina costaricensis), 222 Maclura tinctoria, 326, 327 Macoubea mesoamericana, 66 Macrocnemum glabrescens (Macrocnemum roseum), 398 Macrocnemum pastoense (Macrocnemum roseum), 398 Macrocnemum roseum, 398, 399 madero negro (Gliricidia sepium), 224 madroño (Alibertia edulis), 380 madroño (Amaioua corymbosa), 382 madroño (Calycophyllum candidissimum), 384 madroño (Drypetes standleyi), 374 madroño (Garcinia intermedia), 136 madroño (Garcinia madruno), 136 madroño (Macrocnemum roseum), 398 Magnolia sororum, 268, 269 mahogany (Swietenia macrophylla), 316 majagua (Heliocarpus americanus), 288 majaguillo (Heliocarpus americanus), 288 majaguillo (Muntingia calabura), 334 majaguillo (Trichospermum galeottii), 290 mala sombra (Guapira standleyana), 358 malagueto (Xylopia sericea), 62 malagueto de montaña (Oxandra panamensis), 54 malagueto de montaña (Xylopia macrantha), 62 malagueto hembra (Xylopia aromatica), 60 malagueto macho (Xylopia frutescens), 60 Malmea sp. (Mosannona garwoodii), 52 Malouetia isthmica, 66, 67

malvecino (Acosmium panamense), 214 malvecino (Guapira standleyana), 358 malvecino (Lonchocarpus minimiflorus), 226 malvecino (Swartzia panamensis), 234 mameicillo (Alseis blackiana), 382 mameicillo (Chrysophyllum colombianum), 436 mameicillo (Clethra lanata), 132 mameicillo (Ecclinusa lanceolata), 438 mameicillo (Pouteria glomerata), 442 mameicillo (Sloanea terniflora), 154 mamey (Pouteria fossicola), 440 mamey de montaña (Pouteria fossicola), 440 mamón de montaña (Talisia nervosa), 434 manca caballo (Prosopis juliflora), 210 manga larga (Casearia arborea), 416 manga larga (Laetia procera), 420 mangabé (Pourouma bicolor), 458 mangabé (Schefflera morototoni), 76 Mangifera indica, 34 mangle (Myrsine coriacea), 348 mangle blanco (Laguncularia racemosa), 148 mangle botón (Conocarpus erectus), 146 mangle botoncillo (Conocarpus erectus), 146 mangle cimarrón (Cyrilla racemiflora), 152 mangle colorado (Rhizophora mangle), 380 mangle de montaña (Myrsine coriacea), 348 mangle de sabanas (Myrsine coriacea), 348 mangle negro (Avicennia bicolor), 26 mangle negro (Avicennia germinans), 28 mangle piñuelo (Pelliciera rhizophorae), 362 mangle prieto (Avicennia bicolor), 26 mangle prieto (Avicennia germinans), 28 mangle rojo (Rhizophora mangle), 380 mangle salado (Avicennia bicolor), 26 mangle salado (Avicennia germinans), 28

Index manglillo (Citharexylum caudatum), 460 manglillo (Ternstroemia tepezapote), 452 manglillo de botón (Ternstroemia tepezapote), 452 mango (Mangifera indica), 34 Manihot esculenta, 170 Manilkara bidentata, 438, 439 Manilkara darienensis (Manilkara bidentata), 438 Manilkara meridionalis (Manilkara zapota), 440 Manilkara zapota, 440, 441 manioc (Manihot esculenta), 170 manzanillo de la playa (Hippomane mancinella), 168 manzanillo de sabana (Ternstroemia tepezapote), 452 maquenqué (Oenocarpus mapora), 84 Maquira costaricana (Maquira guianensis), 328 Maquira guianensis, 328, 329 marañón (Anacardium occidentale), 32 marañón curasao (Syzygium malaccense), 356 Maranthes panamensis, 130, 131 maraquita de marea (Amphitecna latifolia), 88 mareon (Bucida buceras), 146 Margaritaria nobilis, 366, 367 maría (Calophyllum longifolium), 132 maría enano (Calophyllum nubicola), 134 Marila laxiflora, 138, 139 Marila pluricostata, 138, 139 mastate (Brosimum utile), 324 mastate blanco (Castilla elastica), 324 mata hombro (Cornus disciflora), 150 mata piojo (Trichilia hirta), 316 mata piojo (Trichilia martiana), 318 mata ratón (Gliricidia sepium), 224 Matayba apetala, 430, 431 Matayba scrobiculata, 432, 433 matillo (Matayba scrobiculata), 432 mauro (Casearia commersoniana), 418 Mayna longicuspis (Mayna odorata), 30 Mayna odorata, 30 mayo (Vochysia ferruginea), 462 membrillo (Gustavia superba), 264 membrillo de montaña (Cespedesia spathulata), 360

membrillo macho (Cespedesia spathulata), 360 mesquite (Prosopis juliflora), 210 Mexican hat (Heisteria concinna), 156 michurago (Ladenbergia macrocarpa), 396 Mickey Mouse (Ouratea lucens), 360 Miconia affinis, 302, 303 Miconia argentea, 304, 305 Miconia centronioides, 304, 305 Miconia elata, 304 Miconia hondurensis, 306, 307 Miconia hyperprasina (Miconia affinis), 302 Miconia impetiolaris, 306, 307 Miconia nervosa, 308, 309 Miconia oinochrophylla, 308 Miconia trinervia, 308, 309 Micropholis melinoniana, 440 Micropholis mexicana (Micropholis melinoniana), 440 miguelario (Compsoneura capitellata), 336 miguelario (Compsoneura sprucei), 338 miguelario (Otoba acuminata), 338 miguelario (Otoba novogranatensis), 340 miguelario (Virola macrocarpa), 342 miguelario (Virola surinamensis), 344 Mimosa tenuiflora, 206, 207 Mimusops darienensis (Manilkara bidentata), 438 Minquartia guianensis, 158 moco de pavo (Citharexylum caudatum), 460 mongo (Crateva tapia), 118 monkey comb (Apeiba membranacea), 286 mora (Maclura tinctoria), 326 Morella cerifera, 336, 337 morillo (Sorocea affinis), 332 morillo (Trophis racemosa), 334 moro (Maclura tinctoria), 326 Mortoniodendron anisophyllum, 296, 297 Mosannona garwoodii, 52, 53 Mosquitoxylum jamaicense, 34, 35 mostrenco (Randia armata), 408 mother-in-law’s lips (Psychotria poeppigiana), 408 Mouriri myrtilloides, 310, 311 Mouriri parvifolia (Mouriri myrtilloides), 310 muñeco (Cordia bicolor), 98 muñeco (Cordia panamensis), 104

489

Muntingia calabura, 334, 335 Myrcia gatunensis, 354, 355 Myrica cerifera (Morella cerifera), 336 Myrica mexicana (Morella cerifera), 336 Myrica xalapensis (Morella cerifera), 336 Myroxylon balsamum, 228, 229 Myrsine coriacea, 348, 349 Myrsine microcalyx (Myrsine coriacea), 348 Myrsine myricoides (Myrsine coriacea), 348 nagua blanca (Alchornea costaricensis), 162 nance (Byrsonima crassifolia), 270 nance (Byrsonima spicata), 272 nance macho (Clethra lanata), 132 nancillo (Byrsonima spicata), 272 nancillo (Clethra lanata), 132 nancito (Byrsonima spicata), 272 nancito (Clethra lanata), 132 naranjita (Swartzia simplex), 236 naranjito (Swartzia simplex), 236 naranjo de monte (Swartzia simplex), 236 Naucleopsis naga, 328, 329 nazareno (Jacaranda copaia), 90 nazareno (Peltogyne purpurea), 182 Nectandra cuspidata 256, 257 Nectandra gentlei (Nectandra cuspidata), 256 Nectandra latifolia (Nectandra purpurea), 258 Nectandra lineata, 256, 257 Nectandra purpurea, 258, 259 Nectandra whitei (Ocotea whitei), 260 Neea amplifolia, 358, 359 Neea elegans (Neea amplifolia), 358 negrito (Annona spraguei), 42 negro (Piptocoma discolor), 86 nené (Ormosia macrocalyx), 230 niguita (Cordia panamensis), 104 nisperillo (Chrysophyllum colombianum), 436 nisperillo (Ecclinusa lanceolata), 438 nisperillo (Pouteria glomerata), 442 níspero (Manilkara bidentata), 438 níspero (Manilkara zapota), 440 níspero de montaña (Manilkara bidentata), 438 noli (Cryosophila warscewiczii), 80 nuno (Hura crepitans), 168

490

Index

Ochroma lagopus (Ochroma pyramidale), 276 Ochroma pyramidale, 276, 277 Ocotea cernua, 258, 259 Ocotea eusericea (Ocotea whitei), 260 Ocotea insularis, 260, 261 Ocotea ira (Ocotea insularis), 260 Ocotea skutchii (Ocotea whitei), 260 Ocotea whitei, 260, 261 Oenocarpus mapora, 84, 85 Oenocarpus panamanus (Oenocarpus mapora), 84 Oleiocarpon panamense (Dipteryx oleifera), 220 olivo (Sapium glandulosum), 172 olivo (Simarouba amara), 448 olla de mono (Lecythis ampla), 266 ollito (Eschweilera pittieri), 264 ollito (Eschweilera sessilis), 264 Olmedia aspera (Trophis caucana), 332 orange (Citrus sinensis), 410 oreja de mula (Miconia impetiolaris), 306 orey (Campnosperma panamense), 34 orinera (Warszewiczia coccinea), 410 Ormosia amazonica, 228, 229 Ormosia coccinea, 230, 231 Ormosia croatii (Ormosia coccinea), 230 Ormosia macrocalyx, 230, 231 Otoba acuminata, 338, 339 Otoba novogranatensis, 340, 341 Ouratea lucens, 360, 361 Oxandra longipetala, 52, 53 Oxandra panamensis, 54, 55 pacaya (Chamaedorea tepejilote), 80 Pachira aquatica, 276, 277 Pachira quinata, 278, 279 Pachira sessilis, 278, 279 pacito (Muntingia calabura), 334 paico (Cordia eriostigma), 102 paico (Cordia lasiocalyx), 102 paico (Cordia porcata), 104 Palicourea guianensis, 398, 399 palito feo (Acalypha diversifolia), 160 palma aceitera (Elaeis oleifera), 82 palma conga (Welfia regia), 84 palma de zancos (Socratea exorrhiza), 84 palma escoba (Cryosophila warscewiczii), 80 palma negra (Astrocaryum standleyanum), 78

palma real (Attalea butyracea), 80 palma tigre (Welfia regia), 84 palo blanco (Laetia thamnia), 422 palo chancho (Hieronyma alchorneoides), 364 palo cuadrado (Macrocnemum roseum), 398 palo de bobo (Erythrina fusca), 222 palo de buba (Jacaranda copaia), 90 palo de chancho (Tetrathylacium johansenii), 424 palo de collar (Ormosia macrocalyx), 230 palo de guaco (Crateva tapia), 118 palo de gusano (Maranthes panamensis), 130 palo de la cruz (Zanthoxylum panamense), 412 palo de pico (Maquira guianensis), 328 palo de pico (Naucleopsis naga), 328 palo de vaca (Brosimum utile), 324 palo del paraíso (Couroupita guianensis), 262 palo santo (Couroupita guianensis), 262 palo santo (Erythrina costaricensis), 222 palo santo (Erythrina fusca), 222 palo santo (Triplaris cumingiana), 372 palo verde (Parkinsonia aculeata), 182 palomo (Aegiphila panamensis), 250 palomo (Citharexylum caudatum), 460 panamá (Sterculia apetala), 294 pantano (Amanoa guianensis), 362 pantano (Tetrathylacium johansenii), 424 papaya (Carica papaya), 118 papayito (Jacaratia spinosa), 118 papelillo (Conostegia xalapensis), 300 papelillo (Miconia argentea), 304 papelillo (Miconia centronioides), 304 papelillo (Miconia elata), 304 papo de montaña (Wercklea cocleana), 292 papo de montaña (Wercklea woodsonii), 294 Parathesis amplifolia, 348, 349 parimontón (Hasseltia floribunda), 420 Parkia nitida, 208, 209 Parkinsonia aculeata, 182, 183 Parmentiera cereifera, 90, 91

pasmo (Siparuna pauciflora), 450 pasmo hediondo (Siparuna pauciflora), 450 pata de elefante (Jacaranda copaia), 90 peine de mono (Apeiba tibourbou), 286 peinecillo (Apeiba membranacea), 286 peinecillo (Apeiba tibourbou), 286 pellejo de gallina (Pera arborea), 172 pellejo de sapo (Licania affinis), 126 pellejo de sapo (Parkia nitida), 208 Pelliciera rhizophorae, 362 Peltogyne purpurea, 182 Peltophorum inerme (Peltophorum pterocarpum), 182 Peltophorum pterocarpum, 182 pene de chombo (Iriartea deltoidea), 82 Pentagonia macrophylla, 400, 401 Pentagonia pubescens (Pentagonia macrophylla), 400 Pera arborea, 172, 173 Perebea costaricana (Maquira guianensis), 328 Perebea xanthochyma, 330, 331 perguetano (Crateva tapia), 118 periquito (Muntingia calabura), 334 peronil (Ormosia amazonica), 228 peronil (Ormosia coccinea), 230 peronil (Ormosia macrocalyx), 230 Persea americana, 260 pezuña de vaca (Cynometra bauhiniifolia), 180 Phoebe cinnamomifolia (Cinnamomum triplinerve), 254 Phoebe insularis (Ocotea insularis), 260 Phoebe johnstonii (Cinnamomum triplinerve), 254 Phoebe mexicana (Cinnamomum triplinerve), 254 pica lengua (Banara guianensis), 414 pica lengua (Casearia arborea), 416 pica lengua (Casearia commersoniana), 418 pica lengua (Casearia sylvestris), 418 pichindé (Zygia longifolia), 214 pico de chombo (Iriartea deltoidea), 82 pico de negro (Iriartea deltoidea), 82 pie de santo (Pelliciera rhizophorae), 362 piedro (Hieronyma alchorneoides), 364

Index pilón (Hieronyma alchorneoides), 364 pimiento (Myrcia gatunensis), 354 pinotea (Podocarpus guatemalensis), 368 pinta mozo (Vismia baccifera), 242 pinta mozo (Vismia billbergiana), 244 pinta mozo (Vismia macrophylla), 244 Piper reticulatum, 366, 367 Piptocoma discolor, 86, 87 Pithecellobium barbourianum (Abarema barbouriana), 186 Pithecellobium chagrense (Zygia latifolia), 212 Pithecellobium latifolium (Zygia latifolia), 212 Pithecellobium longifolium (Zygia longifolia), 214 Pithecellobium oblongum (Pithecellobium unguis­cati), 208 Pithecellobium rufescens (Cojoba rufescens), 188 Pithecellobium saman (Samanea saman), 212 Pithecellobium unguis­cati, 208, 209 pito (Erythrina costaricensis), 222 pito (Erythrina fusca), 222 Pittoniotis trichantha, 400 Platymiscium pinnatum, 232, 233 Platymiscium polystachyum (Platymiscium pinnatum), 232 Platypodium elegans, 232, 233 playo blanco (Hampea appendiculata), 292 Plumeria acutifolia (Plumeria rubra), 68 Plumeria rubra, 68, 69 Pochota quinata (Pachira quinata), 278 Pochota sessilis (Pachira sessilis), 278 Podocarpus guatemalensis, 368, 369 Pogonopus exsertus, 400, 401 Pogonopus speciosus (Pogonopus exsertus), 400 pond apple (Annona glabra), 38 poro-poro (Cochlospermum vitifolium), 144 Posoqueria latifolia, 402, 403 Poulsenia armata, 330, 331 Pourouma aspera (Pourouma bicolor), 458 Pourouma bicolor, 458, 459 Pourouma chocoana (Pourouma bicolor), 458 Pourouma johnstonii (Pourouma bicolor), 458

Pourouma scobina (Pourouma bicolor), 458 Pourouma tessmannii (Pourouma bicolor), 458 Pouteria cooperi (Pouteria torta), 444 Pouteria fossicola, 440, 441 Pouteria glomerata, 442, 443 Pouteria neglecta (Pouteria torta), 444 Pouteria reticulata, 442, 443 Pouteria stipitata, 444, 445 Pouteria torta, 444, 445 Pouteria tuberculata (Pouteria torta), 444 Pouteria unilocularis (Pouteria reticulata), 442 prende-prende (Acalypha diversifolia), 160 Prioria copaifera, 182, 183 Prosopis juliflora, 210, 211 Protium costaricense, 106, 107 Protium panamense, 108, 109 Protium tenuifolium, 108, 109 Pseudobombax septenatum, 280, 281 Pseudosamanea guachapele, 210, 211 Psidium guajava, 356, 357 Psidium guineense, 356, 357 Psychotria elata, 402, 403 Psychotria grandis, 404, 405 Psychotria horizontalis, 404, 405 Psychotria luxurians, 406, 407 Psychotria marginata, 406, 407 Psychotria poeppigiana, 408 Pterocarpus belizensis (Pterocarpus officinalis), 234 Pterocarpus hayesii (Pterocarpus rohrii), 234 Pterocarpus officinalis, 234, 235 Pterocarpus rohrii, 234, 235 purple heart (Peltogyne purpurea), 182 Quararibea asterolepis, 280, 281 Quassia amara, 448, 449 Quercus davidsoniae (Quercus insignis), 238 Quercus insignis, 238, 239 Quercus oocarpa (Quercus insignis), 238 Quercus seibertii (Quercus insignis), 238 quina (Coutarea hexandra), 388 quina (Exostema mexicanum), 388 quira (Andira inermis), 216 quira (Platymiscium pinnatum), 232 quita manteca (Conostegia xalapensis), 300

491

rain tree (Samanea saman),212 ramoncito (Sorocea affinis), 332 Randia armata, 408, 409 Randia panamensis (Randia armata), 408 Rapanea myricoides (Myrsine coriacea), 348 rasca (Licania arborea), 128 rascador (Licania arborea), 128 rascador (Ruprechtia costata), 370 raspa (Licania arborea), 128 raspa lengua (Conostegia xalapensis), 300 raspa lengua (Hasseltia floribunda), 420 ratón (Roupala montana), 372 Rauvolfia littoralis, 68, 69 recadito (Palicourea guianensis), 398 red mangrove (Rhizophora mangle), 380 Rhamnus sharpii, 376, 377 Rheedia acuminata (Garcinia madruno), 136 Rheedia edulis (Garcinia intermedia), 136 Rheedia madruno (Garcinia madruno), 136 Rhizophora mangle, 380, 381 Rhizophora samoensis (Rhizophora mangle), 380 Rinorea sylvatica, 462, 463 roble (Quercus insignis), 238 roble (Tabebuia rosea), 94 roble amarillo (Terminalia amazonia), 148 roble de montaña (Quercus insignis), 238 Rollinia jimenezii (Rollinia mucosa), 54 Rollinia microsepala (Rollinia pittieri), 56 Rollinia mucosa, 54, 55 Rollinia permensis (Rollinia mucosa), 54 Rollinia pittieri, 56, 57 ron-ron (Astronium graveolens), 32 Rondeletia buddleioides (Arachnothryx buddleioides), 384 rosa de monte (Brownea macrophylla), 176 rose-apple (Syzygium malaccense), 356 rosetillo (Randia armata), 408 Roupala montana, 372, 373 rubber (Hevea brasiliensis), 166 Rudgea pittieri, 408, 409

492

Index

Ruprechtia costata, 370, 371 Ruptiliocarpon caracolito, 266, 267 Ryania speciosa, 422, 423 saba (Otoba acuminata), 338 Sacoglottis trichogyna, 240, 241 salamo (Calycophyllum candidissimum), 384 salvia de montaña (Guettarda crispiflora), 394 Samanea saman, 212, 213 sándalo (Myroxylon balsamum), 228 sandbox tree (Hura crepitans), 168 sande (Brosimum utile), 324 sangare (Croton draco), 166 sangre (Licania platypus), 130 sangre (Virola sebifera), 342 sangre de drago (Croton draco), 166 sangre de gallo (Pterocarpus officinalis), 234 sangre de gallo (Pterocarpus rohrii), 234 sangrilillo (Compsoneura capitellata), 336 sangrilillo (Compsoneura sprucei), 338 sangrillo (Bocconia frutescens), 362 sangrillo (Croton billbergianus), 164 sangrillo (Croton draco), 166 sangrillo (Virola koschnyi), 340 sangrillo (Vismia baccifera), 242 sangrillo (Vismia macrophylla), 244 sangrillo de montaña (Otoba acuminata), 338 sangrillo de montaña (Otoba novogranatensis), 340 sanguinaria (Pogonopus exsertus), 400 sanguinaria (Warszewiczia coccinea), 410 santa maría (Calophyllum longifolium), 132 santa maría (Calophyllum nubicola), 134 Sapindus saponaria, 432, 433 sapito (Pera arborea), 172 Sapium aucuparium (Sapium glandulosum), 172 Sapium biglandulosum (Sapium glandulosum), 172 Sapium caudatum (Sapium glandulosum), 172 Sapium glandulosum, 172, 173 Sapium oligoneurum (Sapium glandulosum), 172

sapote (Licania platypus), 130 sapote (Pachira aquatica), 276 sapote negro (Diospyros artanthifolia), 154 sastra (Garcinia intermedia), 136 sastra (Garcinia madruno),136 sastro (Garcinia intermedia), 136 sastro (Garcinia madruno), 136 sauquillo (Hedyosmum bonplandianum), 120 sauquillo (Hedyosmum costaricense), 122 sauquillo (Hedyosmum mexicanum), 122 Scheelea zonensis (Attalea butyracea), 80 Schefflera morototoni, 76, 77 Schizolobium parahyba, 184, 185 Sciadodendron excelsum, 78, 79 sea grape (Coccoloba uvifera), 370 Senna dariensis, 184 Senna reticulata, 184, 185 sequara (Eugenia nesiotica), 352 sequara (Mouriri myrtilloides), 310 Sideroxylon celastrinum, 446, 447 sigua (Cinnamomum triplinerve), 254 sigua (Nectandra cuspidata), 256 sigua (Nectandra lineata), 256 sigua (Nectandra purpurea), 258 sigua (Ocotea cernua), 258 sigua (Ocotea insularis), 260 sigua blanca (Cinnamomum triplinerve), 254 sigua blanco (Caryodaphnopsis burgeri), 254 sigua negro (Guatteria aberrans), 46 sigua negro (Guatteria dumetorum), 48 Simarouba amara, 448, 449 Simarouba glauca (Simarouba amara), 448 Siparuna pauciflora, 450, 451 Sloanea terniflora, 154, 155 soapberry (Sapindus saponaria), 432 Socratea durissima (Socratea exorrhiza), 84 Socratea exorrhiza, 84, 85 solacra (Mouriri myrtilloides), 310 sombrecito (Heisteria acuminata), 156 sombrecito (Heisteria concinna), 156 sombrerito del diablo (Psychotria elata), 402 sombrerito del diablo (Psychotria poeppigiana), 408

Sorocea affinis, 332, 333 Spachea correae, 272 Spachea membranacea, 272, 273 Spanish cedar (Cedrela odorata), 312 Spathodea campanulata, 90 Spondias mombin, 34, 35 Spondias radlkoferi, 36, 37 Stemmadenia grandiflora, 70, 71 Stemmadenia obovata, 70, 71 Sterculia apetala, 294, 295 Stylogyne guatemalensis (Stylogyne turbacensis), 350 Stylogyne nicaraguensis (Stylogyne turbacensis), 350 Stylogyne standleyi (Stylogyne turbacensis), 350 Stylogyne turbacensis, 350, 351 suela (Pterocarpus officinalis), 234 suela (Pterocarpus rohrii), 234 suicide tree (Tachigali versicolor), 186 Swartzia panamensis, 234 Swartzia simplex, 236, 237 Sweetia panamensis (Acosmium panamense), 214 Swietenia macrophylla, 316, 317 Symphonia globulifera, 140, 141 Syzygium malaccense, 356 tabaquillo (Aegiphila anomala), 248 tabaquillo (Aegiphila odontophylla), 250 tabaquillo (Bocconia frutescens), 362 tabaquillo (Cosmibuena macrocarpa), 386 Tabebuia guayacan, 92, 93 Tabebuia ochracea, 92, 93 Tabebuia rosea, 94, 95 Tabernaemontana arborea, 72, 73 Tabernaemontana obliqua (Tabernaemontana undulata), 74 Tabernaemontana panamensis, 72, 73 Tabernaemontana undulata, 74, 75 Tachigali versicolor, 186, 187 tachuelo (Zanthoxylum acuminatum), 412 tachuelo (Zanthoxylum ekmanii), 412 tachuelo (Zanthoxylum panamense), 412 tachuelo (Zanthoxylum setulosum), 414 Talisia croatii, 434 Talisia dwyeri (Talisia nervosa), 434 Talisia nervosa, 434, 435 tangaré (Carapa guianensis), 310

Index Tapirira guianensis, 36, 37 Tapirira myriantha (Tapirira guianensis), 36 tea mangrove (Pelliciera rhizophorae), 362 teak (Tectona grandis), 250 teca (Tectona grandis), 250 tecla (Vochysia ferruginea), 462 Tectona grandis, 250 terciopelo (Sloanea terniflora), 154 terciopelo (Trichilia pallida), 318 Terminalia amazonia, 148, 149 Terminalia chiriquensis (Terminalia oblonga), 150 Terminalia oblonga, 150, 151 Terminalia obovata (Terminalia amazonia), 148 Ternstroemia seemannii (Ternstroemia tepezapote), 452 Ternstroemia tepezapote, 452, 453 teta de chola (Pentagonia macrophylla), 400 Tetragastris panamensis, 110, 111 Tetrathylacium johansenii, 424, 425 Tetrorchidium euryphyllum, 174, 175 Tetrorchidium gorgonae, 174, 175 Theobroma cacao, 284, 285 Thevetia ahouai, 74, 75 tigrillo (Astronium graveolens), 32 tinecú (Schizolobium parahyba), 184 tolerante (Astronium graveolens), 32 toreta (Annona purpurea), 42 torpedo (Beilschmiedia pendula), 252 totumillo (Amphitecna latifolia), 88 totumo (Crescentia cujete), 88 Tovomita longifolia, 140, 141 Tovomita stylosa, 142, 143 Tovomita weddelliana, 142, 143 Tovomitopsis membranifolia (Tovomita stylosa), 142 Trattinnickia aspera, 110, 111 Trema micrantha, 112, 113 tres bocas (Guarea grandifolia), 312 tres chucitos (Randia armata), 408 Trichilia chiriquina (Trichilia martiana), 318 Trichilia cipo (Trichilia tuberculata), 320 Trichilia hirta, 316, 317 Trichilia japurensis (Trichilia pleeana), 320 Trichilia martiana, 318, 319 Trichilia montana (Trichilia pallida), 318

Trichilia pallida, 318, 319 Trichilia pleeana, 320, 321 Trichilia skutchii (Trichilia pallida), 318 Trichilia tuberculata, 320, 321 Trichilia verrucosa (Trichilia pleeana), 320 Trichospermum galeottii, 290, 291 Triplaris cumingiana, 372, 373 trompito (Alibertia edulis), 380 trompito (Hieronyma oblonga), 364 tronador (Hura crepitans), 168 Trophis caucana, 332, 333 Trophis racemosa, 334, 335 trupa (Oenocarpus mapora), 84 tuliviejo (Posoqueria latifolia), 402 Turpinia occidentalis, 450, 451 Ulmus mexicana, 454, 455 Unonopsis panamensis, 56, 57 Unonopsis pittieri, 58, 59 Unonopsis sp. nov., 58, 59 uvero (Coccoloba manzinellensis), 368 uvero (Coccoloba uvifera), 370 uvito (Bactris major), 80 uvito (Coccoloba manzinellensis), 368 uvito (Coccoloba uvifera), 370 uvito (Pourouma bicolor), 458 uvito de parra (Bactris major), 80 uvito de playa (Coccoloba uvifera), 370 Vantanea depleta, 242, 243 Vantanea occidentalis (Vantanea depleta), 242 vaquero (Croton billbergianus), 164 vaquero (Dendropanax arboreus), 76 vara santa (Triplaris cumingiana), 372 Vatairea erythrocarpa, 236, 237 velario (Compsoneura capitellata), 336 velario (Compsoneura sprucei), 338 velario (Otoba acuminata), 338 velario (Otoba novogranatensis), 340 velario (Virola macrocarpa), 342 velario colorado (Virola sebifera), 342 Viburnum costaricanum, 30, 31 Virola koschnyi, 340, 341 Virola macrocarpa, 342, 343 Virola multiflora, 342 Virola nobilis (Virola surinamensis), 344 Virola sebifera, 342, 343 Virola surinamensis, 344, 345 Vismia baccifera, 242, 243

493

Vismia billbergiana, 244, 245 Vismia macrophylla, 244, 245 Vismia panamensis (Vismia baccifera), 242 Vitex cooperi, 252, 253 Vochysia ferruginea, 462, 463 volador (Aspidosperma spruceanum), 64 volador (Gyrocarpus americanus), 238 Warszewiczia coccinea, 410, 411 Welfia georgii (Welfia regia), 84 Welfia regia, 84 Wercklea cocleana, 292, 293 Wercklea woodsonii, 294, 295 white mangrove (Laguncularia racemosa), 148 wild cashew (Anacardium excelsum), 30 wild frangipani (Laxoplumeria tessmannii), 66 wild plumeria (Laxoplumeria tessmannii), 66 Xylopia aromatica, 60, 61 Xylopia bocatorena, 60 Xylopia frutescens, 60, 61 Xylopia longifolia (Xylopia aromatica), 60 Xylopia macrantha, 62, 63 Xylopia sericea, 62, 63 Xylopia xylopioides (Xylopia aromatica), 60 Xylosma chlorantha, 424 yaya (Unonopsis pittieri), 58 yayito (Cymbopetalum lanugipetalum), 44 yayito (Desmopsis panamensis), 44 yayito (Guatteria amplifolia), 48 yayito (Guatteria recurvisepala), 50 yerce (Casearia commersoniana), 418 yuca (Manihot esculenta), 170 yuco de monte (Pachira sessilis), 278 Zanthoxylum acuminatum, 412, 413 Zanthoxylum belizense (Zanthoxylum ekmanii), 412 Zanthoxylum ekmanii, 412, 413 Zanthoxylum jaimei (Zanthoxylum panamense), 412 Zanthoxylum juniperinum (Zanthoxylum acuminatum), 412 Zanthoxylum panamense, 412

494

Index

Zanthoxylum procerum (Zanthoxylum acuminatum), 412 Zanthoxylum setulosum, 414, 415 Zanthoxylum tripetalum (Zanthoxylum acuminatum), 412 zapatero (Hieronyma alchorneoides), 364

zarza (Enterolobium schomburgkii), 190 Ziziphus mauritiana, 378, 379 zopilote (Hernandia didymantha), 238 zorro (Astronium graveolens), 32 zorro (Couratari guianensis), 262

zorro (Lonchocarpus heptaphyllus), 224 Zuelania guidonia, 424, 425 zumbo (Alibertia edulis), 380 Zygia latifolia, 212, 213 Zygia longifolia, 214, 215