Desert Volume 29, Number 2 Published by The University of Arizona for Plants the Boyce Thompson Arboretum A Guide to North American Grasslands David E. Brown and Elizabeth Makings Relict Great Basin Shrub-Grassland near Wupatki National Monument northeast of Flagstaff, Coconino County, Arizona, 1,650 m (5,413 ft). Volume 29, Number 2 Desert Plants Published by The University of Arizona for the Boyce Thompson Arboretum A journal devoted to broadening knowledge of plants 37615 E US Highway 60 indigenous or adapted to arid and sub-arid regions and Superior, AZ 85173 to encouraging the appreciation of these plants. Copyright 2014. The Arizona Board of Regents on Mark D. Siegwarth, editor behalf of The University of Arizona. The Boyce [email protected] Thompson Arboretum at Superior, Arizona, is cooperatively managed by the Boyce Thompson Production Director: Kim Stone Southwestern Arboretum, Inc., The University of Arizona, and Arizona State Parks. Boyce Thompson Arboretum From the editor As Desert Plants begins its 35th year with a new staff, it in editorship comes the opportunity to rethink what Desert seems somewhat appropriate that as we begin a new chap- Plants is and could be. Desert Plants is devoted to broad- ter in the history of Desert Plants, we start with A Guide ening knowledge of plants indigenous or adapted to arid to North American Grasslands by David E. Brown and Eliza- and sub-arid regions and to encouraging the appreciation of beth Makings. Probably one of the most quoted, used and these plants. With such a broad mandate, it is open to vari- reprinted issues of Desert Plants to this day is Volume 4, ous interpretations. We would certainly welcome comments No. 1-4: Biotic Communities of the American Southwest - United from our readers as to what articles they value most or what States and Mexico, by David E. Brown (editor). Not only are they would like to see more of. Another issue confronting we reaching back to our early years for authorship, I believe us is whether we should move to become a peer reviewed we are maintaining the high standards set back then as well. journal. A peer reviewed journal would open us to addi- So, we certainly hope that we are off to a great start, even if tional authors but may eliminate others. So, the fundamental it is a bit late. question is what type of articles you would prefer. We are very excited about several upcoming issues. We We have tried in the past to have some variety in most hope to have one issue dedicated to the 25-year field eval- issues, except for the ones dedicated to a flora or given uation of various exotic tree species at the Mesa Agricul- subject. Typically, we tried to maintain a balance between tural Center in Yuma, Arizona and another issue dedicated Arizona and international articles, technical and non-tech- to a Vascular and Vegetative Flora of the Ironwood Forest nical articles. As Boyce Thompson Arboretum has grown National Monument in Pima and Pinal Counties, Arizona. over the years, we believe we can expand in all areas, but first Other issues will include articles on the 5-year Arizona we would like to hear from you. Hedgehog (Echinocereus triglochidiatus var. arizonicus) salvage Please send your comments to Desert Plants, 37615 E US project, the role of floral scent in host detection for a soli- Highway 60, Superior, Arizona 85173 or DesertPlants@cals. tary specialist bee and an upcoming genus treatment from arizona.edu. Legumes of Arizona to name a few. Again, we hope you enjoy this issue and sorry for the late With so many great issues on the horizon and the change start. Desert Plants Journal is included with Boojum Club, Picket Post Society, and Director’s Circle level memberships at Boyce Thompson Arboretum. One-year subscriptions are also available: individual $20, individual(foreign) $30, institution $50. Back issues are $5 each, plus $3 shipping and handling. Additional copies of Vol. 29(2) are $20, plus $5 shipping and handling. Mailing address: Boyce Thompson Arboretum 37615 E US Highway 60, Superior, AZ 85173 Website: arboretum.ag.arizona.edu E-mail: [email protected] Telephone: 520.689.2723 4 Desert Plants 29(2) 2014 North American Grasslands Contents 5 Acknowledgements 5 Authors’ Preface 6 Introduction 14 Arctic-Boreal Grasslands 16 Rocky Mountain Alpine and Subalpine Grassland 23 Cascade-Sierran Subalpine Grassland 27 Transvolcanic Alpine and Subalpine Grassland 31 Cold Temperate Grasslands 32 Rocky Mountain Montane Grassland 36 Sierran-Cascade Montane Grassland 39 Madrean Montane Grassland 41 Plains Grassland 55 Great Basin Shrub-Grassland 62 Oregonion Coastal Grassland 66 Warm Temperate Grasslands 67 Gulf Coastal Grassland 71 Semidesert Grassland 83 California Valley Grassland 90 Neotropical Savanna Grasslands 92 Central American Savana Grassland 97 Guatemalan-Guerrean Savanna Grassland 100 Campachian-Veracruz Savanna Grassland 103 Tamaulipan Savanna Grassland 106 Sonoran Savanna Grassland 113 Caribbean Savanna Grassland 117 Other Neotropical Savannas 118 Concluding Remarks 119 References 125 Appendix I. Figures. Worldwide grasslands 130 Appendix II. Plant matrix 152 Appenxix III. Animal matrix Above: Trachypogon spicatus Back cover: Semidesert Grassland in the San Simon Valley, just east of the Price Canyon area. August 2010. Photos: Elizabeth Makings 5 Acknowledgements The authors would like to gratefully acknowledge Andy John Taft, John Alcock, Aimee Kessler, Barbara Neuffer, Casillas and Stella Ruth, former office mates of co-author David and Nancy Pearson, Fernando Zuloaga, David Cook, Elizabeth Makings at the Tonto National Forest Supervi- Lee Frataer, Richard E. Brown, Alejandro Velázquez, Raul sor’s Office. Without their expertise and patience teaching Valdez, Bonnie Swarbrick, Joey Carboneaux, D. F. Costello, the basics of ArcMap, the grasslands map would not have R. R. Humphrey, Ray Turner, Kevin Clark, Ron Cole, David been possible. The authors are also indebted to the many Clendenen, Randall Babb, Louella Brown, Steve Gallizioli, contributing photographers who were so generous with Casa Grande Valley Historical Society, Charles H. Lowe, Jr, not only their photos, but also with species-level descrip- and Anibella Flores. tions that many noted in the captions. The photograpers are: Authors’ Preface This monograph is an expanded treatment of the grass- Both plants and animals are therefore of prime consider- lands originally presented in “Biotic Communities of South- ation in the identification and delineation of the grasslands west U.S. and Northwest Mexico” published in Desert Plants featured herein. The authors may take credit for assembling 1982 (1-4). This expansion was prompted by comments what is hoped to be a useful and enduring guide to the grass- received from a number of biologists working in North lands of North America, but the real credit goes to the scien- America’s “grassland biome” who suggested that we include tists this work has referenced – leaders in grassland ecology and compare the Southwest’s grasslands with their area of including giants such as Daniel Axelrod, Michael Barbour, expertise. Ecologists expressing particular interest in such William Billings, Frederic Clements, Rexford Daubenmire, an expansion included Ray Turner, Alberto Burquez, Orie Aldo Leopold, Jerzy Rzedowski, Forrest Shreve, and John Laucks, Miklos Udvardy, and the late Jim Yoakum. Weaver. Their influential theories and pioneering works laid Co-author David Brown has been to each of the grass- the foundation for our current biogeographical view of land communities presented herein from the arctic-subarctic landscapes. to the tropic-neotropic. Biotic communities visited included Those biologists contributing significantly to the inclusion those in the prairie provinces of Canada; the Eastern, Plains, of animal inhabitants within the biotic communities present- Rocky Mountain, Great Basin, Oregonian, and California ed include Russell Davis (mammals), Janet Ruth (grassland regions of the U.S., and in every Mexican state. Tropic- sparrows), R. Roy Johnson (birds), Rich Glinski (raptors) subtropic grasslands and their inhabitants included those and Randal Babb (herpetofauna). observed in Guatemala, Belize, Nicaragua, Honduras and Costa Rica as well as on several Caribbean islands including David E. Brown Cuba. Arizona State University Not all ecologists will agree with the climatic and biotic [email protected] delineations separating the grasslands, correctly pointing out the continuum concept of R. H. Whittaker and others Elizabeth Makings showing that few if any species share identical altitudinal and Arizona State University climatic limitations. Nonetheless, we consider the plant and [email protected] animal assemblages within the various grasslands depicted to not only be varied and large, but to contribute to a better January 2014 understanding of each grassland’s history and evolution. This presentation therefore incorporates the biotic commu- nity concepts originally proposed by H. S. Swarth, Forrest Shreve, and V. E. Shelford; applied by D. I. Rasmussen, and formulated by C. H. Lowe, Jr. 6 Desert Plants 29(2) 2014 North American Grasslands Grassland communities can be found on every continent Introduction and on most oceanic islands. They occupy extensive areas and their regional names may imprint a specific designation to an “Pride of place must certainly go to the entire biotic province. The Eurasian “steppe,” the Hungar- ian “puszta”, the “pampas” of Argentina, the “campos” of Gramineae…the