Flora of the Gran Desierto and Río Colorado Delta (Southwest Center Series) 0816520445, 9780816520442

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Flora of the Gran Desierto and Río Colorado Delta

Richard Stephen Felger

FLORA 0

~he

i ESTEBN MEXICO NOBTHW

THE SOUTHWEST CENTER SERIES

Ignaz Pfefferkorn, Sonora: A Description of the Province Carl Lumholtz, New Trails in Mexico Buford Pickens, I11e Missions of Northern Sonora: A 1935 Field Documentation Gary Paul Nabhan, editor, Counting Sheep: Twenty lVtzys of Seeing Desert Bighorn Eileen Oktavek, Answered Prayer.s: Miracles and Milagros along the Border Curtis M. Hinsley and David R. Wilcox, editors, Frank Hamilton Cushing and the Hemenway Southwestern Archaeological Expedition, 1886-1889, volume 1: The Southwest in the American Imagination: The Writings o(Sylvester Baxte,; 1881-1899 Lawrence J. Taylor and Maeve Hickey, The Road to Mexico Donna J. Guy and Thomas E. Sheridan, editors, Contested Ground: Comparative Frontiers on the Northern and Southern Edges of the Spanish Empire

Julian D. Hayden, The Sierra Pinacate Paul S. Martin, David Yetman, Mark Fishbein, Phil Jenkins, Thomas R. Van Devender, and Rebecca K. Wilson, editors, Gentry's Rfo Mayo Plants: The Tropical Deciduous Forest and Environs of Northwest Mexico.

W J McGee, Trails to Tiburon: The 1894 and 1895 Field Diaries ofW J McGee, transcribed by Hazel Mcfeely Fontana, annotated and with an introduction by Bernard L. Fontana Richard Stephen Felger, Flora of the Gran Desierto and Rfo Colorado of Northwestern Mexico

I

NORTHWESTERN MEXICO ev RICHARD STEPHEN FELGER

The University of Arizona Press © 2000 The Arizona Board of Regents All rights reserved.

(§) This book is printed on acid-free, archival-quality paper. Manufactured in the United States of America. 05 04 03 02 01 00

6 5 4 3 2 1

Library of Congress Cataloging-in-Publication Data Felger, Richard Stephen Flora of the Gran Desierto and Rio Colorado of northwestern Mexico / Richard S. Felger p. cm. - (The Southwest Center series) Includes bibliographical references (p. ) and index. ISBN 0-8165-2044-5 (alk. paper) 1. Desert plants-Mexico-Sonora (State) 2. Desert plants-Sonoran Desert. I. Title. II. Series. QK21 l.F385 1999 581.7'54'097217-dc21 99-6732 CIP British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library.

For Jocelyn M. Wallace (:;' H. B. Wallace

Gntents

ACKNOWLEDGMENTS _ _ _ _ _ _ _ _ _ _ _ _ _ ix ABBREVIATIONS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ xi PART I. THE ENVIRONMENT AND HUMAN INTERACTIONS _ _ _ _ _ _ _ _ _ 3 PALEOCLIMATE _ _ _ _ _ _ _ _ _ _ _ _ _ _ 3 PRESENT CLIMATE _ _ _ _ _ _ _ _ _ _ _ _ _ 6 MAJOR HABITATS _ _ _ _ _ _ _ _ _ _ _ _ _ _ 8

SONOYTA REGION _ _ _ _ _ _ _ _ _ _ _ _ g PINACATE VOLCANIC FIELD _ _ _ _ _ _ _ _ _ 9

Peripheral Lava Flows _ _ _ _ _ _ _ _ 12 The Craters _ _ _ _ _ _ _ _ _ _ _ _ 13 Higher Elevations of the Sierra Pinacate

15

GRANITIC RANGES. _ _ _ _ _ _ _ _ _ _ _ 17 SIERRA DEL ROSARI0 _ _ _ _ _ _ _ _ _ _ 17 DUNES

_ _ _ _ _ _ _ _ _ _ _ _ _ _ 19

DESERT PLAINS OR FLATS_ _ _ _ _ _ _ _ _21 COASTAL HABITATS _ _ _ _ _ _ _ _ _ _ _22 WETLAND HABITATS _ _ _ _ _ _ _ _ _ _23

Tidal Wetlands _ _ _ _ _ _ _ _ _ _ _23 Oases _ _ _ _ _ _ _ _ _ _ _ _ _ _24 Rivers _ _ _ _ _ _ _ _ _ _ _ _ _26 XERORIPARIAN HABITATS _ _ _ _ _ _ _ _ _29 HISTORY AND HUMAN INFLUENCES _ _ _ _ _ _ _ 29

CONSERVATION _ _ _ _ _ _ _ _ _ _ _ _ 31 AGRICULTURE AND HORTICULTURE

_ _ _ _ _ 32

NON-NATIVE PLANTS _ _ _ _ _ _ _ _ _ _ 33

viii CONTENTS

GROWTH FORMS _ _ _ _ _ _ _ _ _ _ _

34

BOTANICAL HISTORY _ _ _ _ _ _ _ _ _ _ _ _ 37 PART

II. THE FLORA - - ~ - - - - - - - _ , 3 9

KEY TO THE MAJOR PLANT GROUPS _ _ _ _ _ _40 PTERIDOPHYTES: FERNS AND FERN ALLIES

42

GYMNOSPERMS: CONE-BEARING PLANTS

49

ANGIOSPERMS-FL OWERING PLANTS: DICOTYLEDONS _ _ _ _ _ _ _ _ _ _ _ _ _ 51 ANGIOSPERMS-FLO WERING PLANTS: MONOCOTYLEDONS _ _ _ _ _ _ _ _ _ _469

GAZETTEER _ _ _ _ _ _ _ _ _ _ _ _ _ _ 565 APPENDIXES A. GROWTH FORMS AND DISTRIBUTION OF PLANTS IN NORTHWESTERN SONORA _ _ _ _ 577 B. DISTRIBUTION OF PLANTS IN SYKES CRATER

595

C. COMMONLY CULTIVATED TREES AND SHRUBS IN NORTHWESTERN SONORA _ _ _ _ _ _ _ _ 599 D. NON-NATIVE PLANTS IN NORTHWESTERN SONORA

603

E. SYSTEMATIC ARRANGEMENT AND RELATIVE ABUNDANCE OF THE GRASSES IN NORTHWESTERN SONORA _ _ _ _ _ _ _ _ _ _ _ _ _ _ 607 F. GEOGRAPHIC DISTRIBUTIONS OF GRASSES IN NORTHWESTERN SONORA _ _ _ _ _ _ _ _ _ 611

LITERATURE CITED _ _ _ _ _ _ _ _ _ _ _ _ 615 INDEX - - - - - - - ~ - - - - - - - - - - 6 5 1 TABLES TABLE 1. SUMMARY OF WEATHER DATA FOR NORTHWESTERN SONORA _ _ _ _ _ _ _ _ _ _ 7 TABLE 2. SUMMARY OF THE MAJOR PLANT GROUPS NORTHWESTERN SONORA _ _ _ _ _ _ _ _ _ 41

Acknowledgments

any friends, colleagues, and organizations helped make this book happen. This project was realized by support from the Wallace Research Foundation and earlier from the Wallace Genetic Foundation. I also acknowledge support from the Sunny Knickerbocker Foundation, the Southwest Parks and Monuments Association, and the U.S. Fish and Wildlife Service. Jane Ivancovich contributed during the early part of the work. Publication funds were provided by the Wallace Research Foundation, Sunny Knickerbocker Foundation, Southwest Center of the University of Arizona, James Aronson, Gil Gillenwater, Jean Russell, James Henrickson, Esther Capin, Council Rock Land and Cattle, Dan Felger, Peggy Hitchcock, Jim Hills, Duncan Laurie, Todd Gillenwater, and Leonard Scheff. A significant experience that set me on the path leading to this volume was gained during a six week period, more than twenty-five years ago, when I camped near Tinaja Tule on the western flank of the Sierra Pinacate with Ann Woodin, her family, and various friends. I am grateful for that privilege. Since then many friends and colleagues have contributed additional help and companionship in the field, valuable information, and reviews. I thank all who have helped with the project, especially the following: Dan Austin (Convolvulaceae); Marc Baker (Cactaceae); Theodore M. Barkley (Asteraceae); Tom Bowen (field work); Bill Broyles (field work, geography, history, and sustained encouragement); Katie Bucher {field work); Sterling Bunnell (field work); Alberto B(uquez (field work, ecology, geography, and biological, historical, scientific, and technical information as well as sustained encouragement); Fernando Chiang-Cabrera (Lycium, Solanaceae, and botanical data); Kim Cliffton (field work); Jane Cole (bibliography); Charles Conner (field work and technical information); Kevin Dahl (field work and manuscript assistance); Tom Daniel (Acanthaceae); Mark A. Dimmitt (field work and botanical and ecological info{mation); Luke Evans (field work and geography); Exequiel Ezcurra (field work, ecological and geographical information, and inspiration); Wayne Ferren (Suaeda, Chenopodiaceae); Mark Fishbein (Apocynaceae, Asclepiadaceae, and other botanical information); Paul Fryxell (Malvaceae); Linnea Gentry (editorial assistance); Judy Gibson (botanical and archival data); Gil Gillenwater, Todd Gillenwater, and Troy Gillenwater (field work); Edward Glenn (field work, ecology of the Rio Colorado delta, and halophytes); Lucretia Brezeale Hamilton (illustrations); Mary Frances Hamilton (archival assistance); Julie Hawkins (Parkinsonia, Fabaceae); Trica Oshant Hawkins (field work); Julian Hayden (field work, geography, history, and encouragement); James Henrickson (Aizoaceae, Amaranthaceae, Chenopodiaceae, Portulacaceae, and many other taxa, manuscript reviews, and sustained inspiration); Wendy Hodgson (Agavaceae); Colin Hughes (Fabaceae); Kevin Horstman (maps); Phil Jenkins (Solanaceae, review of the manuscript, and a great deal of botanical and technical expertise); Matthew B. Johnson (Fabaceae, and illustrations); Peter S. Johnson (map coordinates); Gene Joseph (field work); Dennis Kearns (Cucurbitaceae); Peter Kresan (geology, photographs); Meredith A. Lane (Asteraceae); Linda Leigh (field work and extensive help with the manuscript); Cathy Moser Marlett (illustrations); Angelina Martinez Yrizar (ecology, geography, and technical assistance); Michael McClure (field work); Lucinda McDade (Acanthaceae, extensive manuscript review and botanical information); Reid Moran (Crassulaceae); James D. Morefield (Asteraceae); Gary P. Nabhan (field work, O'odham names); Nancy Nicholson (illustrations); James Norris (field work); David Ortiz Reyna (field work); Bruce Parfitt (Cactaceae and other taxa); William D. Peachey (geology and cactus ecology); Donald Pinkava (Cactaceae and manuscript review);

M

X

ACKNOWLEDGMENTS

Barry Prigge (Loasaceae); Adrianne Rankin (field work, ecological and geographical information); Jon Rebman (Cactaceae); Charlotte Reeder (Poaceae, bibliographic and botanical information); John Reeder (Poaceae, bibliographic and botanical information, and manuscript review); Frances Runyan (illustrations); Sue Rutman (field work and technical information); Dean Saxton (O'odham names); Silke Schneider (field work, encouragement, and tolerance); John Semple (Asteraceae); Richard Spellenberg (Nyctaginaceae); Victor Steinmann (field work, Euphorbiaceae, and botanical information); John Strother (Asteraceae); Scott Sundberg (Asteraceae); Barbara Tellman (field work, photographs}; Laurence Toolin (manuscript review); Dale S. Turner (field work and manuscript assistance); Billie Turner (Asteraceae); Raymond M. Turner (ecological and geographical information, and sharing illustrations); Tom Van Devender (manuscript review, ecological and paleo-botanical information); Warren L. Wagner (Onagraceae); Michael Windham (ferns and fern relatives); James J. White (archival illustrations from Hunt Institute for Botanical Documentation); Anita Williams de Alvarez (field work, history, and geography); George Yatskievych (ferns and fern relatives, Lennoaceae, Orobanchaceae, and extensive botanical information and manuscript review); Allan Zimmerman (field work, Cactaceae and other taxa). I especially thank the curators and staff of the various herbaria who provided facilities, hospitality, information, and loans of specimens. In this regard I thank researchers and staff of the herbaria at the Arizona State University, California Academy of Sciences, Desert Botanical Garden, Jepson Herbarium, Instituto de Biologia de la Universidad Nacional Autonoma de Mexico, Missouri Botanical Garden, Rancho Santa Ana Botanic Garden, San Diego Museum of Natural History, University of Texas, and University of California, Berkeley. A major part of my work was carried out at the University of Arizona herbarium, and I sincerely thank Phil Jenkins, Lucinda McDade and other staff members, volunteers, and herbarium scholars for their generous help. I am grateful to the institutions and presses that have granted permission for use of illustrations, including the Hunt Institute for Botanical Documentation, the Sherman Foundation, Stanford University Press, University of Arizona Press, University of California Press, and University of Washington Press. William K. Hartmann and Fisher Books gave permission to use a photograph (© 1989 Fisher Books) from Desert Heart. The New York Botanical Garden granted permission to reprint illustrations from the lntennountain Flora: Vascular Plants of the lntennountain West, U.S.A., vol. 3 part A,© 1977; vol. 4, © 1984; vol. 3 part B, © 1989; vol. 5, © 1994; vol. 6, © 1997, The New York Botanical Garden. The University of California Press, Berkeley, and the Jepson Herbarium granted permission to use illustrations from The fepson manual: higher plants of California, ]. Hickman (ed.), University of California, Berkeley, © 1993, the Regents of the University California. Stanford University Press gave permission to use illustrations from the Illustrated Flora of the Pacific States, 4 vols., by LeRoy Abrams and Roxana Stinchfield Ferris, © 1960 by the Board of Trustees of the Leland Stanford Junior University. James Henrickson has given unwavering encouragement and an enormous wealth of botanical information, shared numerous illustrations by Bobbi Angell and Felicia Bond, and lent his monumental and extremely useful unpublished Flora of tJ1e Chihuahuan Desert Region (Henrickson & Johnston n.d.). Over the years Charlotte Reeder and John Reeder have been a source of inspiration and knowledge-I have been privileged to gain a deep appreciation of the grasses from these intrepid agrostologists. Michael F. Wilson has been involved in many tasks in the making of this book, and I am grateful for his efforts and dedication. I thank Kirsteen E. Anderson for copy editing, Bill Benoit for expert design and production, and attention to detail, and the many artists who contributed illustrations. Bill Broyles and Joseph C. Wilder of the Southwest Center of the University of Arizona made this book a reality.

Abbreviations

he talent of many illustrators and photographers and the cooperation of several institutions have contributed to the illustrations for this book. Each is identified in individual credit lines below the illustrations as follows:

T

ILLUSTRATORS AB-Alfonso Barbosa AC-Agnes Chase AE-Amy Eisenberg AS-Anthony Salazar BA-Bobbi Angell BT-Barbara Tellman CLV-unable to identify CMM-Cathy Moser Marlett DTM-Daniel Trembly MacDougal EDC-E. D. Church ER-Emily Reed FB-Felicia Bond FLS-Frank Lamson-Scribner FR-Frances Runyan ILW-lra L. Wiggins IS-Isaac Sprague JRJ-Jeanne R. Janish KCH-Kevin C. Horstman KD-Karin Douthit KK-Karen Klitz Ks-Kristina Schierenbeck LAV-Linda Ann Vorobik LBH-Lucretia Brezeale Hamilton LH-Leta Hughey LTD-Lauramay T. Dempster MAD-Mark A. Dimmitt MB-Marlo D. Buchmann MBJ-Matthew B. Johnston MBP-Mary Barnas Pomeroy MD-Maggie Day MDB-M. D. Baker (Miss) MFW-Michael F. Wilson ME-Matilda Essig MK-Margaret Kurzius MWG-Mary Wright Gill NE-Nancy Evans Weaver NLN-Nancy L. Nicholson PB-Phyllis Brick PLK-Peter L. Kresan

xii

ABBREVIATIONS

ILLUSTRATORS (cont.) PM-Paul Mirocha PR-P. Roetter RAJ-Robin A. Jess RHP-Robert H. Peebles RJP-Ruth J. Powell RMT-Raymond M. Turner RSF-Richard S. Felger TH-Thomas Holm VK-Valloo Kapadia WKH-William K. Hartmann WM-William S. Moye Ill

ARCHIVES AND HERBARIA ARIZ-University of Arizona, Tucson ASU-Arizona State University, Tempe CAS-California Academy of Sciences, San Francisco C0TEC0CA (HERM0SILL0)-Comisi6n Tecnico Consultiva para la Determinaci6n Regional de los Coeficientes de Agrostadero COTECOCA (MEXICO CITY)-Comisi6n Tecnico Consultiva para la Determinaci6n Regional de los Coeficientes de Agrostadero DAV-University of California, Davis DES-Desert Botanical Garden, Phoenix DS-Dudley Herbarium of Stanford University, San Francisco ENCB-Escuela Nacional de Ciencias Biol6gicas, lnstituto Politecnico Nacional, Mexico City F-Field Museum of Natural History, Chicago HUNT-Hunt Institute for Botanical Documentation, Carnegie Mellon University. Pittsburgh HUNT/H-US-Hitchcock-Chase Collection of Grass Drawings, Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, indefinite loan from Smithsonian Institution HUNT/us-Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, indefinite loan from Smithsonian Institution HUNT/USDA-United States Department of Agriculture Forest Service Collection, Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh JEPS-Jepson Herbarium, Berkeley MEXU-lnstituto de Biologfa, Universidad Nacional Aut6noma de Mexico, Mexico City MO-Missouri Botanical Garden, Saint Louis 0RPI-Organ Pipe Cactus National Monument, Arizona P0M-Pomona College, Claremont RSA-Rancho Santa Ana Botanic Garden, Claremont so-San Diego Society of Natural History, San Diego SFC-Sherman Foundation Collection, Corona del Mar TEX-University of Texas, Austin UC-University of California, Berkeley UCR-University of California, Riverside UNM-University of New Mexico, Albuquerque us-United States National Herbarium, Smithsonian Institution, Washington, D.C.

Throughout this book, an asterisk(*) denotes a non-native or introduced, exotic plant.

t

indicates a plant that is expected in the Gran Desierto region given its known range, but that has not actually been collected in the region.

FLORA 0

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. i ESTERN MEXICO NQRTl:JW

Part

The Environment and Human Interactions

n a warm spring day the desert breeze is alive with buzzing insects and the smell of yellow palo verde flowers. It has been a long day's hike from our camp at Tinaja del Tule to Pinacate Peak, long because of the distance and so many things to see and record. From the cinder-cone summit I see lava fields, great craters, dunes, distant sawtooth desert mountains, the sea, and the Sierra San Pedro Martir more than a hundred miles across the vanquished delta. I imagine a delta still teeming with life. I see a quiet, primordial empire. Less than three months later, during late May, I return to find most of the waterholes dry-even Tinaja del Tule. There are animal bones where the last water disappeared. The heat-the searing heatthe dust, and the dryness penetrate everything. The wildflowers are gone. Even creosote bush leaves are shriveled, brown, and brittle. The northwestern corner of the state of Sonora, Mexico, makes up the area encompassed in this flora, referred to in the text as "the flora area"; it is a logistically convenient eco-political unit (see Felger & Broyles 1997 and maps). The U.S. border provides the northern boundary, the delta of the Rio Colorado and the Mexican portion of the river the western edge, and the Gulf of California the southern limit. The 112°50' W-113°30' W meridians, which roughly coincide with Mexico Highway 8 from Sonoyta to Puerto Pefiasco, make up the approximate eastern edge of the study area. The area is approximately 15,000 kmZ. It is a substantial portion of the extremely arid center, or heart, of the Sonoran Desert. Within this region there are expansive dune fields, maritime strands, a small river, a once-great river and its delta, tidal wetlands, desert plains, steep granitic mountains, desert oases, and an enormous black and red volcanic field featuring its own mountain, lava flows, cinder cones, and formidable craters. Also included is the Quitobaquito oasis, along the international border but mostly on the Arizona side. Despite this aridity the vegetation is surprisingly diverse (Ezcurra 1984; Ezcurra, Equihua, & L6pezPortillo 1987; Felger 1980) . As would be expected of an arid region, however, the flora is relatively small in relation to area (McLaughlin & Bowers 1999). The flora includes 589 species in 327 genera and 85 families . Among these are 79 species not native to the region, although the majority are urban and agricultural weeds that have not become established in undisturbed natural desert habitats.

O

PALEOCLIMATE Wonderfully detailed reconstructions of the vegetation and climate of many desert areas in northern Mexico and the southwestern United States are available for the past 30,000 to 45,000 years (Betancourt, Van Devender, & Martin 1990). These reconstructions are based on analysis of plant contents from fossil packrat middens. The vegetation history of the past several million years is dominated by the advent and conclusion of glacial periods. Past glacial periods have lasted five to ten times as long as interglacials (Porter 1989; Winograd et al. 1997). Thus, climate and vegetation similar to what is in the flora area today would have been present for perhaps only 5% to 10% of the last 1.8 million years. In the flora area ice-age climates have favored the expansion of woodland and chaparral species from higher to lower elevations and from the north to the south. Likewise, species with subtropical affini-

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