
TEXAS TECH UNIVERSITY 

Natural Science Research Laboratory 

Special Publications 

Museum of Texas Tech University 

Number 65 15 December 2016 

Contributions in Natural History: 

A Memorial Volume in Honor of Clyde Jones 


Edited by 

Richard W. Manning, Jim R. Goetze, and Franklin D. Yancey, II 


Artist’s Statement and Memory 

My name is Margaret Goetze. I have a B FA in studio art from Texas A&M Corpus Christi. It was my privilege to 
know Dr. Clyde Jones for almost 20 years, and, from our first associations, I always felt welcomed and accepted into his 
‘circle of scientists.’ Clyde always expressed an interest in my art and it seemed to me that he often was my biggest fan. 
Therefore, I was excited and honored to be commissioned to design the covers for this book, and it took me some time and 
a great deal of thought to complete the commission. 

Front cover .—Anyone who knew or worked with Clyde knows this chair and what it represents. My hardest decision 
was to leave the background blank. This was intentional. Clyde was cosmopolitan in his travels and his research interests 
and, even in the time that I knew him, Clyde and his chair had been many places. Therefore, as you look at the cover, the 
chair and Clyde are wherever you want them to be. 


Back cover. —“Clyde, here is the Spotted Bat, Euderma maculatum, that I always wanted to draw for you. I hope that 
you would be pleased.” 


Special Publications 

Museum of Texas Tech University 

Number 65 


Contributions in Natural History: 

A Memorial Volume in Honor of Clyde Jones 


EDITED BY 


Richard W. Manning, Jim R. Goetze, and Franklin D. Yancey, II 


Layout and Design: Lisa Bradley 
Cover Designs: Margaret Goetze 
Production Editor: Lisa Bradley 

Copyright 2016, Museum of Texas Tech University 

This publication is available free of charge in PDF format from the website of the Natural Sciences Research 
Laboratory, Museum of Texas Tech University (nsrl.ttu.edu). The authors and the Museum of Texas Tech 
University hereby grant permission to interested parties to download or print this publication for personal 
or educational (not for profit) use. Re-publication of any part of this paper in other works is not permitted 
without prior written permission of the Museum of Texas Tech University. 

This book was set in Times New Roman and printed on acid-free paper that meets the guidelines for per¬ 
manence and durability of the Committee on Production Guidelines for Book Longevity of the Council on 
Library Resources. 

Printed: 15 December 2016 


Library of Congress Cataloging-in-Publication Data 

Special Publications of the Museum of Texas Tech University, Number 65 
Series Editor: Robert D. Bradley 

Contributions in Natural History: A Memorial Volume in Honor of Clyde Jones 
Richard W. Manning, Jim R. Goetze, Franklin D. Yancey, II (editors) 


ISSN 0149-1768 
ISBN 1-929330-31-6 
ISBN13 978-1-929330-31-7 


Museum of Texas Tech University 
Lubbock, TX 79409-3191 USA 
(806)742-2442 


Table of Contents 


Editors’ Preface and Acknowledgments 1 

Abbreviated Resume 3 

Bibliography 5 

Prologue: “We learned about field work, and for some reason we loved it”—Oral History 13 

Excerpts from the Reminiscences of Clyde Jones 

David Marshall 

A New Species of Myotis (Chiroptera: Vespertilionidae) from Suriname 49 

Ricardo Moratelli, Don E. Wilson, Alfred L. Gardner, Robert D. Fisher, and 
EliecerE. Gutierrez 

Non-volant Mammals of Ash Meadows National Wildlife Refuge, Nevada 67 

Richard W. Manning and Martin R. Heaney 

A Resurvey of Bats at Dinosaur National Monument 97 

Michael A. Bogan and Tony R. Mollhagen 

Distribution Records and Reported Sightings of the White-nosed Coati (. Nasua narica ) in 127 

Texas, with Comments on the Species’ Population and Conservation Status 

David J. Schmidly, John Karges, and Robert Dean 

Ecological Distribution and Foraging Activity of the Ghost-faced Bat {Mormoops 147 

megalophylla) in Big Bend Ranch State Park, Texas 

Franklin D. Yancey, II 

Pleistocene/Holocene Faunas from the Trans-Pecos 157 

Arthur H. Harris 

First Documented Record of Nutting’s Flycatcher ( Myiarchus nuttingi) for Texas 177 

Mark W. Lockwood 

Bats of Kimball and Cheyenne Counties in the Panhandle of Nebraska 183 

Kenneth N. Geluso and Keith Geluso 

Molecular Systematics and Phylogeny of Peromyscus nudipes (Cricetidae: Neotominae) 201 

Robert D. Bradley, Maria Nuhez-Tabares, Taylor J. Soniat, Sara Kerr, 

Russell W. Raymond, and Nicte Ordohez-Garza 

An Inventory of Bats in Arch Canyon, San Juan County, Utah 215 

Tony R. Mollhagen and Michael A. Bogan 


225 


Comparison of Pasturelands Containing Texas Kangaroo Rat (Dipodomys elator ) Burrows 
to Adjacent Roadsides in Wichita County, Texas, with Comments on Road Usage 
by D. elator 

Jim R. Goetze, Allan D. Nelson, and Larry L. Choate 


Encomia and Reflections 

233 

Photographs 

259 


Editors’ Preface and Acknowledgments 


It is with great pleasure that we present this volume containing research works, encomia, photographs, 
and art work in honor of Clyde Jones. Clyde was a teacher, mentor, colleague, and dear friend, as well as doc¬ 
toral committee chair or co-chair for the three of us. Discussion of a volume in honor of Clyde Jones first was 
entertained in June 2013, in a group setting between the three of us and Mark Lockwood, Mike Bogan, Cindy 
Ramotnik, and Mary Ann Jones. This occurred during a field trip at Big Bend Ranch State Park in the Trans-Pecos 
region of Texas. Clyde participated in the trip, but was unaware of our conversations regarding the planning of 
this volume. We originally discussed possible publication of a festschrift honoring Clyde, but unfortunately he 
passed away on 6 April 2015 after a lengthy period of illness. 

In July 2015, we decided to proceed with work on a memorial volume to honor him. We began inquiries 
regarding the possibility of publication of such a book through the Museum of Texas Tech University. We then 
solicited potential contributors—a group that included former colleagues and graduate students of Clyde’s, and 
other individuals who had been associated with or worked with Clyde throughout his long and distinguished career. 

Although not everyone contacted was able to respond and contribute to this volume, nonetheless this work 
contains 11 research articles ranging in scope from Pleistocene/Holocene faunal analyses, ecological and ethologi- 
cal studies on terrestrial and volant mammals, the description of a new species of Myotis , and resurrection and 
redefinition of an ‘older’ species of Peromyscus, to a first record of an avian species in Texas, and even a rather 
botanically-related research study. If you consider the wide-ranging and eclectic research interests of Clyde 
Jones, as evidenced in the included Abbreviated Resume and Bibliography, we think that “El Jefe” (as we often 
referred to CJ) would be pleased with the contributions. 

We also decided to include encomia or “songs of praise” in honor of Dr. Jones, memory statements, a 
transcript of an interview with Clyde, and photographs. We believe that all of the aforementioned works and 
contributions will allow readers a glimpse of the complex character, breadth of knowledge, and outstanding ac¬ 
complishments of Clyde Jones. Some of the materials may, hopefully, even make readers smile, and we are quite 
sure that Clyde would enjoy that! 

This project required tremendous contributions of time and effort from Robert D. Bradley, Series Editor, and 
Lisa Bradley, Production Editor, of the Natural Science Research Laboratory (NSRL) of the Museum of Texas 
Tech University. In addition, many external reviewers generously gave their time and provided valuable sugges¬ 
tions for the papers. We thank Fred Stangl and Terry Maxwell for providing final reviews of the entire volume. 

In addition to publishing this volume, much of the cost of production was subsidized by the NSRL of the 
Museum of Texas Tech University. We wish to thank the NSRL, as well as the Texas Tech University Libraries 
and several individuals, for their generous contributions toward the production of this volume. 

Clyde could exhibit a ‘stormy’ personality at times (at least when necessary in order to accomplish a task or 
properly ‘motivate’ an individual) but, more often, he displayed an open, generous, friendly temperament and a 
warm smile for those that he met. Dr. Jones often portrayed himself as a simple country boy from the Nebraska 
Sandhills (see Resume and Prologue) and indeed he was that person. However, as those who knew him can attest 
and those who read this volume will learn, Clyde Jones was much, much more... 

Richard W. Manning 
Jim R. Goetze 
Franklin D. Yancey, II 

1 


Clyde Jones 

1935-2015 


Abbreviated Resume 


General Information .—Born in Scottsbluff, Nebraska, 3 March 1935; married; two married children. 

Education .—Burwell High School, Burwell, Nebraska, graduated 1952; Hastings College, Hastings, Ne¬ 
braska, graduated 1957 with BA in History, Biology, and Education; University of New Mexico, Albuquerque, 
New Mexico, graduated 1960 with MS in Zoology and Botany; University of New Mexico, Albuquerque, New 
Mexico, graduated in 1964 with PhD in Biology. 

Professional Employment History. —University of New Mexico, Department of Biology: Field Research 
Assistant, 1958-1959; Graduate Assistant, 1957-1961; Teaching Assistant, 1961-1962; Assistant Curator, Museum 
of Southwestern Biology, 1962-1964. Tulane University, Department of Biology: Assistant Professor of Biology, 
1965-1969. Delta Regional Primate Research Center, Covington, Louisiana: Research Associate, 1967-1969. 
United States Fish and Wildlife Service: National Museum of Natural History, Chief of Mammal Section, Bird 
and Mammal Laboratories, 1970-1973; Director of National Fish and Wildlife Laboratory, 1973-1979; Denver 
Wildlife Research Center, Director, 1979-1982. Texas Tech University: Director of the Museum, Texas Tech 
University, 1982-1985; Department of Museum Science, Chairman, 1982-1987; Department of Biological Sci¬ 
ences, Professor, 1987-1999, Paul Whitfield Horn Professor, 1999-2003, Paul Whitfield Horn Professor Emeritus, 
2004-2015. 

Professional Organizations .—American Society of Mammalogists (life member); Asociacion Mexicana De 
Mastozoologia, A.C.; Asociacion Mexicana Para El Estudio De Los Mamiferos Marinos, A.C.; Big Bend Natural 
History Association; Biological Society of Washington; Chihuahuan Desert Research Institute; Society for Big 
Bend Studies; Society of Systematic Biologists; Southwestern Association of Naturalists; Texas Academy of 
Science; Texas Society of Mammalogists; Washington Biologist’s Field Club. 

Professional Organization Activities .—American Society of Mammalogists (various committee member¬ 
ship, Board of Directors, Editor for Reviews, Managing Editor); Asociacion Mexicana Para El Estudio De Los 
Mamiferos Marinos, A.C. Vocale (served on committees, as an elected officer, as an editor, managing editor, and 
board member); Biological Society of Washington (council member, Vice President, President); Chihuahuan Desert 
Research Institute (Board of Scientists member); Texas Society of Mammalogists (served on various committees, 
established scholarship fund, President Elect, President, Executive Committee). 

Task Forces and Additional Committee Service .—Antarctic Observer; Antarctic Inspection Team; Whale 
Policy Committee; International Conference on the Biology of Whales; Marine Mammal Task Force; Antarctic 
Task Force; Research Management System Development Task Force; Delegation to the Plenipotentiary Confer¬ 
ence (CITES); Lacey Act Proposed Interpretation Task Force; Antarctic Resources Study Panel and Sub-Panel; 
U.S. and Mexico Joint Commission on Wildlife Conservation; Permits Study Committee; Kangaroo Manage¬ 
ment Evaluation Committee; Joint NOAA-FWS International Marine Mammal Task Force; U.S. and Mexico 
Joint Committee on Marine Mammals; Wildlife Biology Advisory Committee; Wild Boar Management Advisory 
Committee; CSRS Review Team; Rosenthal Seminar Series; Interagency Grizzly Bear Technical Review Panel; 
Joint Council on Food and Agricultural Sciences; Interagency Grizzly Bear Steering Committee; Old Growth as 
Wildlife Habitat Program; Museum Assessment Program; Museum Studies Committee; Zuni Museum Advisory 
Board. Also served on numerous committees within the Texas Tech University System. 

Teaching Experience .—Hastings High School (Biology), 1956; Hasting College (Invertebrate Zoology 
laboratory), 1956; University of New Mexico (laboratories in General Biology, General Zoology, Plant Anatomy, 


3 


4 


Clyde Jones Memorial Volume 


Cytology, and Histology), 1957-1962; Tulane University (lectures and laboratories in Terrestrial Ecology, General 
Biology, Environmental Biology, Mammalogy, Comparative Anatomy, Biology of Primates), 1965-1966, 1969; 
University of Maryland (lectures and labs in Mammalogy, team taught), 1975; Colorado State University (lectures 
and labs in Vertebrate Pest Management, team taught), 1981; Texas Tech University (lectures in Professional 
Ethics in Museums, Collection Management, Zoological Nomenclature, Field Methods, Biology of Animals, 
Vertebrate Structure and Development, Introduction to Mammalogy, Advanced Mammalogy, Special Topics in 
Mammalogy, Vertebrate Natural History, Bat Communities, History of Mammalogy, Scholarly Writing in Zool¬ 
ogy, and graduate labs in Field Methods), 1984-2002. 

Awards .—Phi Sigma Outstanding Graduate Student Award; USFWS Outstanding Performance Award; 
Antarctic Service Medal; USFWS Special Achievement Award; Commendation from the Secretary, U.S. De¬ 
partment of the Interior; USFWS Letter of Commendation; USFWS Quality Performance Award; Texas Tech 
University Press Award of Excellence; Outstanding Researcher Award Texas Tech University; American Society 
of Mammalogists Hartley H. T. Jackson Award; Paul Whitfield Horn Professor Award, Texas Tech University; 
Paul Whitfield Horn Professor Emeritus Award, Texas Tech University. 

Honors .—Phi Sigma; Sigma Xi; Phi Kappa Phi; American Men and Women of Science; Research Associ¬ 
ate Department of Vertebrate Zoology National Museum of Natural History; Research Associate Museum of 
Southwestern Biology; Senior Executive Service (Charter Member) U.S. Civil Service Commission; Associate 
of International Center for Arid and Semi-arid Land Studies; Athletic Hall of Fame, Hastings College; Fellow of 
Texas Academy of Science; Honorary Member Texas Society of Mammalogists; Honorary Member American 
Society of Mammalogists. 

Graduate Students .—Served on the following graduate student’s committees either as chairperson or co-chair 
for Masters or PhD degree. Tulane University: Merilyn Warkentin Hasler (MS), Fernando Alvarez (PhD), John 
Pagels (PhD), Frances Miller Cashner (PhD). Texas Tech University: Stephen MacDonald (MA), Mary Candee 
(MA), Pat Brown (MA), Tommy Eaton (MA), Lorelei Mount (MA), Brenda Cooke (MA), Nancy Hildreth (MA), 
Patsy Jackson (MA), Mark Murphy (MA), David Zuflacht (MA), Dawn Kaufman (MS), Deidre Parish (MS), 
Maryann Lynch (MS), Kristie Jo Roberts (MS), Robert Hollander (PhD), Paisley Cato (PhD), Richard Manning 
(PhD), Larry Choate (PhD), Jim Goetze (PhD), Franklin Yancey, II (PhD). 

Research Interests .—Taxonomy, systematics, distribution, ecology, and biogeography of Recent mammals, 
especially bats and rodents of America, as well as bats, primates, and rodents of West Africa. Distribution, ecol¬ 
ogy and status of amphibians and reptiles in North America. Biodiversity, conservation, and management of 
mammals and wildlife habitats. Management and conservation of specimens in museums. 


Editors’note: For an excellent, abbreviated autobiography, refer to Dr. Jones’ article entitled “You Have to Catch 
Them First” that is found within the book Going Afield (2005), published by the Museum of Texas Tech Univer¬ 
sity, Lubbock (Carleton J. Phillips and Clyde Jones, editors). A PDF copy may be obtained at the following URL 
address: http://www.nsrl.ttu.edu/publications/otherpubs/Going%20Afield.pdf. 


Bibliography 


200. Yancey, F. D., II, R. W. Manning, J. R. Goetze, L. L. 
Lindsey, R. D. Bradley, and C. Jones. The hooded skunk 
(Mephitis macroura) from the Davis Mountains of West 
Texas, with comments on the species’ natural history, 
morphology, molecular characteristics, and conservation 
status. Texas Journal of Science, in review. 

199. Jones, C., F. D. Yancey, II, and R. W. Manning. 2015. 
The mammals of Caprock Canyons State Park, Texas. 
Occasional Papers, Museum of Texas Tech University 
335:1-26. 

198. Bradley, R. D., L. K. Ammerman, R. J. Baker, L. C. Brad¬ 
ley, J. A. Cook, R. C. Dowler, C. Jones, D. J. Schmidly, 
F. B. Stangl, Jr., R. A. Van Den Bussche, and B. Wursig. 
2014. Revised checklist of North American mammals 
north of Mexico, 2014. Occasional Papers, Museum of 
Texas Tech University 327:1-27. 

197. Suttkus, R. D., and C. Jones. 2013. Fishes of Devils 
River, Val Verde County, Texas. Occasional Papers, 
Museum of Texas Tech University 317:1-8. 

196. Jones, C., M. W. Lockwood, T. R. Mollhagen, F. D. 
Yancey, II, and M. A. Bogan. 2011. Mammals of the 
Chinati Mountains State Natural Area, Texas. Occasional 
Papers, Museum of Texas Tech University 300:1-29. 

195. Jones, C., and M. W. Lockwood. 2008. Additions to the 
mammalian fauna of Big Bend Ranch State Park, Texas. 
Occasional Papers, Museum of Texas Tech University 
282:1-3. 

194. Manning, R. W., C. Jones, and F. D. Yancey, II. 2008. 
Annotated checklist of Recent land mammals of Texas, 
2008. Occasional Papers, Museum of Texas Tech Uni¬ 
versity 278:1-18. 

193. Baker, R. J., C. Jones, R. E. Martin, and L. C. Bradley. 
2007. History of the Texas Society of Mammalogists. 
Special Publications, Museum of Texas Tech University 
52:1-60. 

192. Haynie, M. L., J. G. Brant, L. Rex McAliley, J. P. Car¬ 
rera, M. A. Revelez, D. A Parish, X. Viteri, C. Jones, and 
C. J. Phillips. 2006. Investigations of a natural corridor 
between two national parks in central Ecuador: Results 
from the Sowell Expedition, 2001. Occasional Papers, 
Museum of Texas Tech University 263:1-16. 

191. Manning, R. W., F. D. Yancey, II, and C. Jones. 2006. 
Morphometric variation in two populations of the cactus 
mouse ( Peromyscus eremicus ) from Trans-Pecos Texas. 
Occasional Papers, Museum of Texas Tech University 
262:1-5. 

190. Yancey, F. D., II, and C. Jones. 2006. Changes of dis¬ 
tributions of bats in Texas. Occasional Papers, Museum 
of Texas Tech University 258:1-5. 


189. Yancey, F. D., II, R. W. Manning, and C. Jones. 2006. 
Mammals of the Harte Ranch area of Big Bend National 
Park, Brewster County, Texas. Occasional Papers, Mu¬ 
seum of Texas Tech University 253:1-15. 

188. Kennedy, S., and C. Jones. 2006. Two new records of 
mammals from the Davis Mountains, Jeff Davis County, 
Texas. Occasional Papers, Museum of Texas Tech Uni¬ 
versity 252:1-^1. 

187. Suttkus R. D., and C. Jones. 2006. Fishes of Inde¬ 
pendence Creek and Pecos River. Occasional Papers, 
Museum of Texas Tech University 248:1-7. 

186. Wickliffe, J. K., R. D. Bradley, F. B. Stangl, Jr., J. L. Pat¬ 
ton, D. A Parish, C. Jones, D. J. Schmidly, and R. J. Baker. 
2005. Molecular systematics and phylogeographic his¬ 
tory of Thomomys bottae in Texas. Pp. 507-522 in Contri- 
buciones mastozoologicas, en homenaje a Bernardo Villa 
(V. Sanchez-Cordero and R. A Medellin, eds.). Instituto 
de Biologia, UNAM, and Instituto de Ecologia, UNAM, 
Conabio, Mexico. 706 pp. 

185. Suttkus, R. D., and C. Jones. 2005. Atlas of fishes of 
the upper Red River system in Texas and Oklahoma. 
Occasional Papers, Museum of Texas Tech University 
246:1-32. 

184. Brant, J. G., and C. Jones. 2005. Annotated checklist of 
marine mammals of Texas. Occasional Papers, Museum 
of Texas Tech University 244:1-4. 

183. Bradley, L. C., B. R. Amman, J. G. Brant, R. McAliley, 
F. Mendez-Harclerode, J. R. Suchecki, C. Jones, H. H. 
Genoways, R. J. Baker, and R. D. Bradley. 2005. Mam¬ 
malogy at Texas Tech University: A historical perspective. 
Occasional Papers, Museum of Texas Tech University 
243:1-30. 

182. Jones, C. 2005. You have to catch them first. Pp. 185— 
199 in Going afield (C. J. Phillips and C. Jones, eds.). 
Museum of Texas Tech University, Lubbock. 289 pp. 

181. Stangl, Jr., F. B., M. M. Shipley, J. R. Goetze, and C. 
Jones. 2005. Comments on the predator-prey relation¬ 
ship of the Texas kangaroo rat (Dipodomys elator ) and 
barn owl (Tyto alba). American Midland Naturalist 153: 
135-141. 

180. DeBaca, R. S., and C. Jones. 2004. Report on the mam¬ 
mals of the Davis Mountains: An emphasis on the species 
documented at Texas Nature Conservancy properties. 
Report to the Nature Conservancy of Texas. 355 pp. 

179. Baker, R. J., L. C. Bradley, R. D. Bradley, J. W. Dragoo, 
M. D. Engstrom, R. S. Hoffmann, C. A. Jones, F. Reid, D. 
W. Rice, and C. Jones. 2003. Revised checklist of North 
American mammals north of Mexico, 2003. Occasional 
Papers, Museum of Texas Tech University 229:1-23. 


5 


6 


Clyde Jones Memorial Volume 


178. Brant, J. G., J. L. Higginbotham, and C. Jones. 2002. 
Noteworthy records of the silver-haired bat, Lasionycte- 
ris noctiagans (Chiroptera:Vespertilionidae) in Presidio 
County, Texas. Southwestern Naturalist 47:633-635. 

177. Higginbotham, J. L., R. S. DeBaca, J. G. Brant, and 
C. Jones. 2002. Noteworthy records of bats from the 
Trans-Pecos region of Texas. Texas Journal of Science 
54:277-282. 

176. Brant, J. G., and C. Jones. 2002. Distributional records of 
ma mm als from the Permian Basin, Texas. Texas Journal 
of Science 54:269-276. 

175. DeBaca, R. S., and C. Jones. 2002. The ghost-faced bat, 
Mormoops megalophila (Chiroptera: Mormoopidae), 
from the Davis Mountains, Texas. Texas Journal of 
Science 54:89-91. 

174. Carroll, D. S., L. L. Peppers, R. D. Bradley, and C. Jones. 

2002. Sigmodon ochrognathus is a monotypic species: 

Evidence from DNA sequences. Southwestern Naturalist 
47:494-497. 

173. Frey, J. K., R. D. Fisher, M.A. Bogan, and C. Jones. 2002. 

F irst record of the Arizona cotton rat ( Sigmodon arizonae ) 
in New Mexico. Southwestern Naturalist 47:491-493. 

172. Jones, C. 2002. Afterword. Pp. 469 in Texas natural his¬ 
tory: A century of change (D. J. Schmidly, author). Texas 
Tech University Press, Fubbock. 534 pp. 

171. Schmidly, D. J., and C. Jones. 2001. 20th Century 
changes in mammals and mammalian habitats along the 
Rio Grande/Rio Bravo from Fort Quitman to Amistad. 

Pp. 177-204 in Binational Symposium Rio Grande/Rio 
Bravo: Ft. Quitman to Amistad Reservoir. Proceedings, 

14 June 2000. Cuidad Juarez, Chihuahua, Mexico. U.S. 
Department of the Interior, Secretariat of Environment 
and Natural Resources, International Boundary and Water 
Commission, Washington, D.C. x + 301 pp. 

170. Higginbotham, J. F., and C. Jones. 2001. The southeast¬ 
ern myotis, Myotis austroriparius (Chiroptera: Vespertil- 
ionidae), from Comanche County, Texas. Texas Journal 
of Science 53:193-195. 

169. Jones, C., and D. A. Parish. 2001. Effects of the Pecos 
River on the geographic distributions of mammals in 
western Texas. Occasional Papers, Museum of Texas 
Tech University 204:1-11. 

168. Edwards, C. W., D. S. Carroll, M. F. Clary, K. E. 
Halcomb, M. F. Haynie, S. R. Hoofer, F. G. Hoffmann, 

M. B. O’Neill, E. Webb, M. J. Hamilton, R. A. Van Den 
Bussche, D. J. Schmidly, C. Jones, and R. D. Bradley. 

2000. Records of mammals from northeast and south 
Texas. Occasional Papers, Museum of Texas Tech 
University 200:1-8. 

167. Mantooth, S. J., C. Jones, and R. D. Bradley. 2000. 
Molecular systematics of Dipodomys elator (Rodentia: 
Heteromyidae) and its phylogeographic implications. 155. 
Journal of Mammalogy 81:885-894. 


166. Fockwood, M. W., and C. Jones. 2000. Vertebrate 
remains found in barn owl pellets from Crosby County, 
Texas. Texas Journal of Science 52:169-173. 

165. Jones, C. 2000. Vertebrate predators on bats in North 
America north of Mexico. Pp. 229-241 in Reflections 
of a naturalist: Papers honoring Professor Eugene D. 
Fleharty (J. R. Choate, ed.). Fort Hays Studies, Special 
Issue 1:1-241. 

164. Yancey, F. D., II, and C. Jones. 2000. Distribution and 
ecologic relationships of pocket mice ( Chaetodipus ) in the 
Big Bend region of Texas. Occasional Papers, Museum 
of Texas Tech University 195:1-14. 

163. Bradley, R. D., R. J. Baker, C. Jones, N. C. Parker, D. J. 
Schmidly, Y. Ackerson, D. H. Riskind, and R. R. George. 
2000. Faunal surveys of state-owned properties. P. 53 in 
1999 Wildlife research highlights (R. C. Telfair, II, ed.). 
Texas Parks and Wildlife Department, Austin, 41-60. 

162. Suttkus, R. D., and C. Jones. 1999. Observations on the 
nine-banded armadillo, Dasypus novemcinctus, in south¬ 
ern Fouisiana. Tulane Studies in Zoology and Botany 
31:1-22. 

161. Jones, C., and R. D. Bradley. 1999. Notes on red bats, 
Lasiurus (Chiroptera: Vespertilionidae), of the Davis 
Mountains and vicinity, Texas. Texas Journal of Science 
51:341-344. 

160. Bradley, R. D., D. S. Carroll, M. F. Clary, C. W. Ed¬ 
wards, I. Tiemann-Boege, M. J. Hamilton, R. A. Van Den 
Bussche, and C. Jones. 1999. Comments on some small 
mammals from the Big Bend and Trans-Pecos regions 
of Texas. Occasional Papers, Museum of Texas Tech 
University 193:1-6. 

159. Yancey, F. D., II, and C. Jones. 1999. Collared pika 
Ochotona collaris. Pp. 677-678 in The Smithsonian book 
of North American mammals (D. E. Wilson and S. Ruff, 
eds.). Smithsonian Institution Press, Washington, D C. 
xxv + 750 pp. 

158. Yancey, F. D., II, and C. Jones. 1999. Pinyon mouse 
Peromyscus truei. Pp. 582-583 in The Smithsonian book 
of North American mammals (D. E. Wilson and S. Ruff, 
eds.). Smithsonian Institution Press, Washington, D.C., 
xxv + 750 pp. 

157. Wallace, A. M., and C. Jones. 1999. Stephen’s woodrat 
Neotoma stephensi. P. 611 in The Smithsonian book of 
North American mammals (D. E. Wilson and S. Ruff, 
eds.). Smithsonian Institution Press, Washington, D.C., 
xxv + 750 pp. 

156. Parish, D. A., and C. Jones. 1999. Big free-tailed bat 
Nyctinomops macrotis. Pp. 130-131 in The Smithsonian 
book of North American mammals (D. E. Wilson and S. 
Ruff, eds.). Smithsonian Institution Press, Washington, 
D.C., xxv + 750 pp. 

Mauk-Cunningham, C., and C. Jones. 1999. Southeastern 
myotis Myotis austroriparius. Pp. 83-85 in The Smithson- 


Bibliography 


7 


ian book of North American mammals (D. E. Wilson and 
S. Ruff. eds.). Smithsonian Institution Press, Washington, 
D C., xxv + 750 pp. 

154. Lynch, M. R., and C. Jones. 1999. Rafinesque’s big- 
eared bat Corynorhinus rafinesquii. Pp. 119-121 in The 
Smithsonian book of North American mammals (D. E. 
Wilson and S. Ruff, eds.). Smithsonian Institution Press, 
Washington, D.C. xxv + 750 pp. 

153. Goetze, J. R., and C. Jones. 1999. Texas kangaroo rat 
Dipodomys elator. Pp. 527-528 in The Smithsonian book 
of North American mammals (D. E. Wilson and S. Ruff, 
eds.). Smithsonian Institution Press, Washington, D.C., 
xxv + 750 pp. 

152. Gharaibeh, B. M., and C. Jones. 1999. Hooded skunk 
Mephitis macroura. Pp. 186-187 in The Smithsonian 
book of North American mammals (D. E. Wilson and S. 
Ruff, eds.). Smithsonian Institution Press, Washington, 
D.C., xxv + 750 pp. 

151. Yancey, F. D., II, and C. Jones. 1999. Alopecia in the 
white-ankled mouse, Peromyscus pectoralis (Mam¬ 
malia: Rodentia), in Texas. Texas Journal of Science 
51:271-272. 

150. Jones, C., L. Hedges, and K. Bryan. 1999. The western 
yellow bat, Lasiurus xanthinus (Chiroptera: Vespertilioni- 
dae), from the Davis Mountains, Texas. Texas Journal of 
Science 51:267-269. 

149. Bradley, R. D., D. J. Schmidly, and C. Jones. 1999. The 
northern rock mouse, Peromyscus nasutus (Mammalia: 
Rodentia), from the Davis Mountains, Texas. Occasional 
Papers, Museum of Texas Tech University 190:1-3. 

148. Baker, R. J., C. J. Phillips, R. D. Bradley, J. M. Burns, D. 
Cooke, G. F. Edson, O. R. Haragan, C. Jones, R. R. Monk, 
J. T. Montford, D. J. Schmidly, and N. C. Parker. 1998. 
Bioinformatics, museums, and society: Integrating bio¬ 
logical data for knowledge-based decisions. Occasional 
Papers, Museum of Texas Tech University 187:1-4. 

147. Parker, N. C., R. D. Bradley, J. M. Burns, G. F. Edson, 
D. R. Haragan, C. Jones, R. R. Monk, J. T. Montford, 
C. J. Phillips, D. J. Schmidly, and R. J. Baker. 1998. 
Bio informatics: A multidisciplinary approach for the 
life sciences. Occasional Papers, Museum of Texas Tech 
University 186:1-8. 

146. Manning, R. W., and C. Jones. 1998. Annotated check¬ 
list of Recent land mammals of Texas, 1998. Occasional 
Papers, Museum of Texas Tech University 182:1-20. 

145. Choate, L. L., and C. Jones. 1998. Annotated checklist of 
Recent land mammals of Oklahoma. Occasional Papers, 
Museum of Texas Tech University 181:1-13. 

144. Yancey, F. D., II, J. R. Goetze, and C. Jones. 1998. Sac- 
copteryx leptura. Mammalian Species 582:1-3. 

143. Yancey, F. D., II, J. R. Goetze, and C. Jones. 1998. Sac- 
copteryx bilineata. Mammalian Species 581:1-5. 


142. Yancey, F. D., II, R. W. Manning, J. R. Goetze, and C. 
Jones. 1998. The mammals of Lake Meredith National 
Recreation Area and adjacent areas, Hutchinson, Moore, 
and Potter counties, Texas. Occasional Papers, Museum 
of Texas Tech University 174:1-20. 

141. Jones, C., R. S. Hoffmann, D. W. Rice, M. D. Engstrom, 
R. D. Bradley, D. J. Schmidly, C. A. Jones, and R. J. Baker. 
1997. Revised checklist of North American mammals 
north of Mexico. Occasional Papers, Museum of Texas 
Tech University 173:1-20. 

140. Roberts, K. J., F. D. Yancey, II, and C. Jones. 1997. Pre¬ 
dation by great-horned owls on Brazilian free-tailed bats 
in North Texas. Texas Journal of Science 49:215-218. 

139. Parker, N, C., R. J. Baker, R. D. Bradley, C. Jones, R. R. 
Monk, D. J. Schmidly, R. W. Sims, and F. D. Yancey, II. 
1997. Texas Tech Museum and Texas GAP Program: A 
partnership providing field data for GAP analysis. Elec¬ 
tronic publication, http://www. tcru. ttu.edu/tcrux. 11 pp. 

138. Baker, R. J., B. Albin, R. D. Bradley, J. J. Bull, J. M. 
Burns, K. A. Clark, G. Edson, R. E. Estrada, E. Farley, 
C. B. Fedler, B. M. Gharaibeh, R. L. Hammer, C. Jones, 
R. R. Monk, J. T. Montford, G. Moore, N. C. Parker, J. 
Rawlings, A. Sansom, D. J. Schmidly, R. W. Sims, H. 
A. Wichman, and F. D. Yancey, II. 1997. Natural sci¬ 
ence database: Resource management and public health. 
Pp. 10-20 in Collaboration: The key to success (M. 
Shaughnessy, ed.). Proceedings 4th Annual Conference, 
Organization of Fish and Wildlife Information Managers, 
Key Largo, Florida. 96 pp. 

137. Yancey, F. D., II, and C. Jones. 1997. Rafinesque’s 
big-eared bat, Plecotus rafinesquii (Chiroptera: Vesper- 
tilionidae), from Shelby County, Texas. Texas Journal of 
Science 49:166-167. 

136. Yancey, F. D., II, and C. Jones. 1997. Dispersal of two 
species of harvest mice (. Reithrodontomys ) between the 
High Plains and the Rolling Plains of Texas. Occasional 
Papers, Museum of Texas Tech University 166:1-5. 

135. Yancey, F. D., II, P. Raj, S. U. Neill, and C. Jones. 1997. 
Survey of rabies among free-flying bats from the Big 
Bend region of Texas. Occasional Papers, Museum of 
Texas Tech University 165:1-5. 

134. Yancey, F. D., II, W. Meinzer, and C. Jones. 1997. Aber¬ 
rant morphology in western diamondback rattlesnakes 
(Crotalus atrox ). Occasional Papers, Museum of Texas 
Tech University 164:1-4. 

133. Roberts, K. J., F. D. Yancey, II, and C. Jones. 1997. 
Distributional records of small mammals from the Texas 
Panhandle. Texas Journal of Science 49:57-64. 

132. Jones, C., and C. G. Schmitt. 1997. Mammal species 
of concern in New Mexico. Pp. 179-205 in Life among 
the muses: Papers in honor of James S. Findley (T. L. 
Yates, W. L. Gannon, and D. E. Wilson, eds.). Museum 


8 


Clyde Jones Memorial Volume 


of Southwestern Biology, The University of New Mexico, 
Albuquerque. 290 pp. 

131. Yancey, F. D., II, K. J. Roberts, and C. Jones. 1996. 
Prairie falcon predation on Brazilian free-tailed bats. 
Prairie Naturalist 28:146. 

130. Hrachovy, S. K., R. D. Bradley, and C. Jones. 1996. 
Neotoma goldmani. Mammalian Species 545:1-3. 

129. Yates, T. L., C. Jones, and J. A. Cook. 1996. Preserva¬ 
tion of voucher specimens. Pp. 265-273 in Measuring 
and monitoring biological diversity standard methods 
for mammals (D. E. Wilson, F. R. Cole, J. D. Nichols, R. 
Rudran, and M. S. Foster, eds.). Smithsonian Institution 
Press, Washington, D.C. xxvii + 409 pp. 

128. Jones, C., W. J. McShea, M. J. Conroy, and T. H. Kunz. 
1996. Capturing mammals. Pp. 115-155 in Measuring 
and monitoring biological diversity standard methods 
for mammals (D. E. Wilson, F. R. Cole, J. D. Nichols, R. 
Rudran, and M. S. Foster, eds.). Smithsonian Institution 
Press, Washington, D.C. xxvii + 409 pp. 

127. Yancey, F. D., II, and C. Jones. 1996. Notes on three 
species of small mammals from the Big Bend region of 
Texas. Texas Journal of Science 48:247-250. 

126. Manning, R. W., F. D. Yancey, II, and C. Jones. 1996. 
Nongeographic variation and natural history of two 
sympatric species of pocket mice, Chaetodipus nelsoni 
and Chaetodipus eremicus, from Brewster County, Texas. 
Pp. 191-195 in Contributions in mammalogy: A memorial 
volume honoring Dr. J. Knox Jones, Jr. (H. H. Genoways 
and R. J. Baker, eds). Museum of Texas Tech University, 
Lubbock, ii + 315 pp. 

125. Yancey, F. D., II, J. R. Goetze, C. Jones, and R. W. 
Manning. 1996. The mammals of Justiceburg Wildlife 
Management Area and adjacent areas, Garza and Kent 
counties, Texas. Occasional Papers, Museum of Texas 
Tech University 161:1-26. 

124. Goetze, J. R., R. W. Manning, F. D. Yancey, II, and C. 
Jones. 1996. The mammals of Kimble County, Texas. 
Occasional Papers, Museum of Texas Tech University 
160:1-31. 

123. Yancey, F. D., II, and C. Jones. 1996. New county records 
for ten species of bats (Vespertilionidae and Molossidae) 
from Texas. Texas Journal of Science 48:137-142. 

122. C. Jones, C. A. Jones, J. K. Jones, Jr., and D. E. Wilson. 
1996. Pan troglodytes. Mammalian Species 529:1-9. 

121. Gharaibeh, B. M., and C. Jones. 1996. Myosciurus 
pumilio. Mammalian Species 523:1-3. 

120. Findley, J. S., C. Jones, H. H. Genoways, E. C. Birney, 
and R. J. Baker. 1996. Obituary, J. Knox Jones, Jr.: 
1929-1962. Journal of Mammalogy 77:578-593. 

119. Yancey, F. D., II, R. W. Manning, and C. Jones. 1996. 
Distribution, natural history, and status of the Palo Duro 


mouse, Peromyscus truei comanche, in Texas. Texas 
Journal of Science 48:3-12. 

118. Jones, C., and R. W. Manning. 1996. The mammals. 
Pp. 29-38 in Rangeland wildlife (P. R. Krausman, ed.). 
Society for Range Management, Denver, Colorado, xi + 
440 pp. 

117. Yancey, F. D., II, C. Jones, and J. R. Goetze. 1995 . Notes 
on harvest mice ( Reithrodontomys ) of the Big Bend region 
of Texas. Texas Journal of Science 47:263-268. 

116. Manning, R. W., C. Jones, and F. D. Yancey, II. 1995. 
Noteworthy records of amphibians and reptiles from 
northwestern and western Texas. Texas Journal of Science 
47:231-235. 

115. Yancey, F. D., II, C. Jones, and R. W. Manning. 1995. 
The eastern pipistrelle, Pipistrellus subflavus (Chiroptera: 
Vespertilionidae), from the Big Bend region of Texas. 
Texas. Journal of Science 47:229-231. 

114. Yancey, F. D., II, J. R. Goetze, B. M. Gharaibeh, and C. 
Jones. 1995. Distributional records of small mammals 
from the southwestern rolling plains of Texas. Texas 
Journal of Science 47:101-105. 

113. Goetze, J. R., F. D. Yancey, II, C. Jones, and B. M. 
Gharaibeh. 1995. Noteworthy records of mammals from 
the Edwards Plateau of central Texas. Texas Journal of 
Science 47:3-8. 

112. Choate, J. R, J. K. Jones, Jr., and C. Jones. 1994. Hand¬ 
book of mammals of the south-central states. Louisiana 
State University Press, Baton Rouge, ix + 304 pp. 

111. Williams, S. L., A. M. Wallace, and C. Jones. 1993. 
Effect of relative humidity on cranial dimensions of 
mammals. Collection Forum 9:40-46. 

110. Jones, J. K, Jr., R. W. Manning, F. D. Yancey, II, and 
C. Jones. 1993. Records of five species of small mam¬ 
mals from western Texas. Texas Journal of Science 
45:104-105. 

109. Choate, L. L., R. W. Manning, J. K. Jones, Jr., C. Jones, 
and S. Henke. 1992. Mammals from the southern 
border of the Kansan Biotic Province in western Texas. 
Occasional Papers, Museum of Texas Tech University 
152: 1-34. 

108. Jones, C., C. A. Jones, and J. A. Gore. 1992. Eastern 
chipmunk. Pp. 294-299 in Rare and endangered biota of 
Florida, Vol. 1, Mammals (S. R. Humphrey, ed.). Univer¬ 
sity Press Florida, Gainesville, xxviii + 392 pp. 

107. Jones, J.K., Jr., and C. Jones. 1992. Revised checklist of 
Recent land mammals of Texas, with annotations. Texas 
Journal of Science 44:53-74. 

106. Jones, J. K., Jr., R. F. Hoffmann, D. W, Rice, C. Jones, R. 
J. Baker, and M. D. Engstrom. 1992. Revised checklist 
of North American mammals north of Mexico, 1991. 
Occasional Papers, Museum of Texas Tech University 
146:1-23. 


Bibliography 


9 


105. Jones, C., and J. R. Goetze. 1991. Vertebrate remains 
found in barn owl pellets from Wilbarger County, Texas. 
Texas Journal of Science 43:326-328. 

104. Cato, P. S., and C. Jones. 1991. Natural history museum s : 
Directions for growth. Texas Tech University Press, Lub¬ 
bock. iv + 252 pp. 

103. Suttkus, R. D., and C. Jones. 1991. Observations on 
winter and spring reproduction in Peromyscus leucopus 
(Rodentia Muridae) in southern Louisiana. Texas Journal 
of Science 43:179-189. 

102. Choate, L. L, R. W. Manning, J. K. Jones, Jr., C. Jones, 
and T. R. Mollhagen. 1991. Records of mammals from 
the Llano Estacado and adjacent areas of Texas and New 
Mexico. Occasional Papers, Museum of Texas Tech Uni¬ 
versity 138:1-11. 

101. Jones, C., and J.K. Jones, Jr. 1990. Review of Sealander, 
J. A., and G. A. Heidt, Arkansas mammals, their natural 
history, classification, and distribution. University Ar¬ 
kansas Press, Fayetteville, xiv + 308 pp., 1990. Journal 
of Mammalogy 1:705. 

100. Jones, C. 1990. Review of Seal, U. S., E. T. Thome, M. 
A. Bogan, and S. H. Anderson (eds.), Conservation biol¬ 
ogy and the black-footed ferret. Yale University Press, 
New Haven, Connecticut, xvii + 302 pp., 1989. Journal 
of Mammalogy 71:704. 

99. Jones, C. A., and C. Jones. 1990. Review of Kirkland, 
G. L., Jr., and J. N. Layne (eds.) Advances in the study 
of Peromyscus (Rodentia). Texas Tech University Press, 
Lubbock. 366 pp., 1989. Journal of Mammalogy 71:482. 

98. Milner, J., C. Jones, and J. K. Jones, Jr. 1990. Nyctino- 
mops macrotus. Mammalian Species 351:1-4. 

97. Pesaturo, R. J., J. K. Jones, Jr., R. W. Manning, and C. 
Jones. 1990. Mammals of the Muleshoe Sandhills, Bai¬ 
ley, Hale, and Lamb counties, Texas. Occasional Papers, 
Museum of Texas Tech University 136:1-32. 

96. Choate, L. L., J. K. Jones, Jr., R. W. Manning, and C. 
Jones. 1990. Westward ho: Continued dispersal of the 
pygmy mouse, Baiomys taylori, on the Llano Estacado 
and in adjacent areas of Texas. Occasional Papers, Mu¬ 
seum of Texas Tech University 134:1-8. 

95. Hollander, R. R., C. Jones, J. K. Jones, Jr., and R. W. 
Manning. 1990. Preliminary analysis of the effects of the 
Pecos River on geographic distribution of small mammals 
in western Texas. Journal of Big Bend Studies 2:97-107. 

94. Best,T. L.,N. J. Hildreth, and C. Jones. 1989. Dipodomys 
deserti. Mammalian Species 339:1-8. 

93. Jones, C., and C. H. Carter. 1989. Annotated checklist of 
the Recent mammals of Mississippi. Occasional Papers, 
Museum of Texas Tech University 128:1-9. 

92. McCullough, D., and C. Jones. 1989. Review of Zeveloff, 
S. I., Mammals of the Intermountain West. University 


Utah Press, Salt Lake City, xxiv + 365 pp., 1988. Journal 
of Mammalogy 70:453-454. 

91. Jones, C., and R. W. Manning. 1989. Myotis austrori- 
parius. Mammalian Species 332:1-3. 

90. Jones, C., and N. J. Hildreth. 1989. Neotoma stephensi. 
Mammalian Species 328:1-3. 

89. Manning, R. W., J. K. Jones, Jr., and C. Jones. 1989. 
Comments on distribution and variation in the big brown 
bat, Eptesicus fuscus , in Texas. Texas Journal of Science 
41:95-101. 

88. Jones, J. K., Jr., R. W. Manning, C. Jones, and R. R. Hol¬ 
lander. 1988. Mammals of the northern Texas Panhandle. 
Occasional Papers, Museum of Texas Tech University 
126:1-54. 

87. Jones, C. 1988. Review of VanBlaricom, G. R., and 
J.A. Estes (eds.), The community ecology of sea otters. 
Springer-Verlag, Inc., New York, xv + 247 pp., 1988. 
Journal of Mammalogy 69:870-879. 

86. Jones, C., M. A. Bogan, and L. M. Mount. 1988. Status 
of the Texas kangaroo rat ( Dipodomys elator ). Texas 
Journal of Science 40:249-258. 

85. Jones, J. K., Jr., C. Jones, and D. J. Schmidly. 1988. 
Annotated checklist of Recent land mammals of Texas. 
Occasional Papers, Museum of Texas Tech University 
119:1-26. 

84. Manning, R. W., C. Jones, J. K. Jones, Jr., and R. R. 
Hollander. 1988. Subspecific status of the pallid bat, 
Antrozous pallidus, in the Texas Panhandle and adjacent 
areas. Occasional Papers, Museum of Texas Tech Uni¬ 
versity 117:1-5. 

83. Manning, R. W., J. K. Jones, Jr., C. Jones, and R. R. Hol¬ 
lander. 1988. An unusual number of fetuses in the pallid 
bat. Prairie Naturalist 19:261. 

82. Jones, C., R. D. Suttkus, and M. A. Bogan. 1987. Notes 
on some mammals of north-central Texas. Occasional 
Papers, Museum of Texas Tech University 115: 1-21. 

81. Genoways, H. H, C. Jones, and O. L. Rossolimo (eds.). 
1987. Mammal collection management. Texas Tech 
University Press, Lubbock. 219 pp. 

80. Jones, J. K., Jr., R. W. Manning, R. R. Hollander, and C. 
Jones. 1987. Annotated checklist of Recent mammals 
of northwestern Texas. Occasional Papers, Museum of 
Texas Tech University 111: 1-14. 

79. Hollander, R. R., C. Jones, R. W. Manning, and J. K. 
Jones, Jr. 1987. Distributional notes on some mammals 
from the Edwards Plateau and adjacent areas of south- 
central Texas. Occasional Papers, Museum of Texas Tech 
University 110:1-10. 

78. Manning, R. W., J. K. Jones, Jr., R. R. Hollander, and C. 
Jones. 1987. Notes on distribution and natural history of 


10 


Clyde Jones Memorial Volume 


some bats on the Edwards Plateau and in adjacent areas 
of Texas. Texas Journal of Science 39:279-285. 

77. Jones, C., and J. K. Jones, Jr. 1987. Review of Hoff- 
meister, D. F., Mammals of Arizona. University Arizona 
Press, Arizona Game and Fish Dept., Tucson, xx + 602 
pp., 1986. Journal of Mammalogy 68:458-460. 

76. MacDonald, S. O., and C. Jones. 1987. Ochotona col- 
laris. Mammalian Species 281:1-4. 

75. Jones, C. 1987. Review of Goodall, J., The chimpanzees 
of Gombe, patterns of behavior. Belknap Press, Harvard 
University Press, Cambridge, Massachusetts, xiv + 673 
pp., 1986. Journal of Mammalogy 68:201-202. 

74. Jones, C., R. R. Hollander, J. K. Jones, Jr., and R. W. 
Manning. 1987. Noteworthy records of mammals from 
the Texas Panhandle. Texas Journal of Science 39:97-102. 

73. Jones, J. K., Jr., D. C. Carter, H. H. Genoways, R. S. Hoff¬ 
mann, D. W. Rice, and C. Jones. 1986. Revised checklist 
of North American mammals north of Mexico, 1986. 
Occasional Papers, Museum of Texas Tech University 
107:1-22. 

72. Jones, C. 1986. Review of Jones, J. K., Jr., D. M. Arm¬ 
strong, and J. R. Choate, Guide to Mammals of the Plains 
States. University Nebraska Press, Lincoln, xvii + 371 
pp., 1986. Journal of Mammalogy 67:614. 

71. Carter, D. C., W. D. Webster, J. K. Jones, Jr., C. Jones, 
andR. D. Suttkus. 1985. Dipodomys elator. Mammalian 
Species 232:1-3. 

70. Jones, C. 1984. Review of Oxnard, C., The order of 
man: A biomathematical anatomy of the primates. Yale 
University Press, New Haven, Connecticut, xiv + 366 
pp., 1984. Journal of Mammalogy 65:534-535. 

69. Jones, C., and C. A. Jones. 1984. Review of Schmidly, 
D. J., Texas mammals east of the Balcones Fault Zone. 
Texas A&M University Press, College Station, xvii + 400 
pp.,1983. Journal of Mammalogy 65:365-367. 

68. Jones, C. 1984. Tubulidentates, proboscideans, and 
hyracoideans . Pp. 523-535 in Orders and families of 
Recent mammals of the World (S. Anderson and J. K. 
Jones, Jr., eds.). John Wiley and Sons, New York, xii + 

686 pp. 

67. Jones, C. 1984. Review of Beck. B., and C. Wemmer 
(eds.), The biology and management of an extinct species, 
Pere David’s Deer. Noyes Publications, Park Ridge, New 
Jersey xiv +193 pp., 1983. Journal of Mammalogy 65: 
176. 

66. Jones, C. 1983. Review of Woloszyn, D., and B. W. 
Woloszyn, Los mamiferos de la Sierra de La Laguna, Baja 
California Sur. ConsejoNacional de Cienciay Tecnologia, 
Circuito Cultural Universitario, Mexico, D. F. 168 pp., 
1982. Journal of Mammalogy 64:728-729. 

65. Jones, C. 1983. Review of Thornback, J., and M. 
Jenkins (compilers), The IUCN red data book: Part I. 


ICUN, Gland, Switzerland, xl +515 pp., 1982. Journal 
of Mammalogy 64:547-549. 

64. Jones, J. K., Jr., D. M. Armstrong, R. S. Hoffmann, and 

C. Jones. 1983. Mammals of the Northern Great Plains. 
University Nebraska Press, Lincoln, xii + 379 pp. 

63. Jones, C., G. Clemmer, R. D. Suttkus, and R. Curnow. 
1982. Distributional checklist of the mammals along the 
Colorado River in the Grand Canyon. Occasional Papers, 
Tulane University Museum of Natural History 3:1-15. 

62. Jones, C. 1982. Professional ethics and motives. Pp. 
1-4 in The Natural Resource Agency—Its people and or¬ 
ganization (S. Smith and A. Rosenthal, eds.). Occasional 
Papers, U.S. Fish and Wildlife Service, Washington, D.C. 
vi + 16 pp. 

61. Jones, C. 1981. Feeding habits. Pp. 515-516 in World¬ 
wide Furbearer Conference Proceedings (J. Chapman and 

D. Pursely, eds.). Worldwide Furbearer Conference, Inc., 
Frostburg, Maryland, vol. 1, xvii + 651 pp. 

60. Jones, C. 1980. Wildlife research and ecological prob¬ 
lems in Baja California. Pp. 128-137 in Proceedings of 
the National Audubon Society Symposium, The birds of 
Mexico: Their ecology and conservation (P. Schaeffer 
and S. Ehlers, eds.). National Audubon Society Western 
Education Center, Tiburon, California. 137 pp. 

59. Loveless, C. M., J. E. Crawford, C. Jones, R. Linn, T. Ri¬ 
pley, and D. R. Smith. 1979. The role of federal agencies 
in fish and wildlife research. Pp. 188-196 in Transactions 
of the Forty-fourth North American Wildlife and Natural 
Resources Conference (K. Sabol, ed.). Wildlife Manage¬ 
ment Institute, Washington, D.C. 630 pp. 

58. Jones, C., andR. D. Suttkus. 1979. The distribution and 
taxonomy of Tamias striatus at the southern limits of its 
geographic range. Proceedings of the Biological Society 
of Washington 91:828-839. 

57. Jones, C. 1978. Eastern chipmunk, Tamias striatus 
(Linnaeus). Pp. 35-36 in Rare and endangered biota 
of Florida, vol. 1, Mammals (J. N. Layne, ed.). State 
of Florida Game and Fresh Water Fish Commission, 
Tallahassee. xx+ 52 pp. (Published previously in 1976: 
Inventory of rare and endangered biota of Florida, Florida 
Audubon Society and Florida Defenders of the Environ¬ 
ment, Microfiche, 17 cards.) 

56. Jones, C. 1978. Dendrohyrax dorsalis. Mammalian 
Species 113:1-4. 

55. Jones, C., and S. Anderson. 1978. Callicebus moloch. 
Mammalian Species 112:1-5. 

54. Suttkus, R. D., G. H. Clemmer, and C. Jones. 1978. 
Mammals of the riparian region of the Colorado River in 
the Grand Canyon area of Arizona. Occasional Papers, 
Tulane University Museum of Natural History 2:1-23. 

53. Jones, C. 1977. Plecotus rafinesquii. Mammalian Spe¬ 
cies 69:1^1. 


Bibliography 


11 


52. Jones, C. 1976. Economics and conservation. Pp. 
133-145 in Biology of bats of the New World family 
Phyllostomatidae, Part I (R. J. Baker, J. K. Jones, Jr., and 
D. Carter, eds.). Special Publications, Museum of Texas 
Tech University 10:1-218. 

51. Scott, N. J., D. E. Wilson, C. Jones, and R. M. Andrews. 
1976. The choice of perch dimensions by lizards of the 
genus Anolis (Reptillia, Lacertilia, Iguanidae). Journal 
of Herpetology 10:75-84. 

50. Jones, C., and R. D. Suttkus. 1975. Notes on the natural 
history of Plecotus rafinesquii. Occasional Papers of the 
Museum of Zoology, Louisiana State University 47:1-14. 

49. Bogan, M. A.,andC. Jones. 1975. Observations on Lepus 
callotis in New Mexico. Proceedings of the Biological 
Society of Washington 88:45-50. 

48. Findley, J. S., A. H. Harris, D. E. Wilson, and C. Jones. 
1975. Mammals of New Mexico. University New Mexico 
Press, Albuquerque, xxii + 360 pp. 

47. Pagels, J. F., and C. Jones. 1974. Growth and develop¬ 
ment of the free-tailed bat, Tadarida brasilliensis cyno- 
cephala (Le Conte). Southwestern Naturalist 19:267-276. 

46. Riopelle, A., and C. Jones. 1974. Field studies of primates 
in Rio Muni, West Africa, 1967-1968. Pp. 255-262 
in National Geographic Society research reports, 1967 
projects (P Oesher, ed.). National Geographic Society, 
Washington, D.C. x + 323 pp. 

45. Suttkus, R. D., and C. Jones. 1973. Colony structure 
and organization of Pipistrellus subflavus in southern 
Louisiana. Journal of Mammalogy 54:962-968. 

44. Jones, C., and R. D. Fisher. 1973. Comments on the type 
specimen of Neotoma desertorum sola Merriam 1894 
(Mammalia: Rodentia). Proceedings of the Biological 
Society of Washington 86:435-438. 

43. Riopelle, A., and C. Jones. 1973. Field studies of pri¬ 
mates in Rio Muni, West Africa. Pp. 219-223 in National 
Geographic Society research reports, 1967 projects (P. 
Oesher, ed.). National Geographic Society, Washington, 
D.C. x + 323 pp. 

42. Jones, C. 1973. Body temperatures of a Manis gigantean 
and Manis tricuspis. Journal of Mammalogy 54:263-266. 

41. Manville, R, and C. Jones. 1972. Life spans: Mam¬ 
mals. Pp. 229-233 in Biology data book (P Altman and 
D. Dittmer, eds.). Federation of American Society for 
Experimental Biology, Bethesda, Maryland, 2 nd ed., vol. 

1. 606 pp. 

40. Jones, C. 1972. Comparative ecology of three pteropid 
bats in Rio Muni, West Africa. Journal of Zoology, Lon¬ 
don 167:353-370. 

39. Jones, C. 1972. Observations on dental deposits and 
deficiencies of wild talapoin monkeys ( Cercopithecus 
talapoin) collected in Rio Muni, West Africa. Laboratory 
Primate News 11:28-34. 


38. Jones, C. 1972. Natural diets of wild primates. Pp. 
56-77 in Pathology of simian primates, Part I: General 
pathology (R. N. T.-W. Fiennes, ed.). S. Karger, Basel, 
Switzerland. 929 pp. 

37. Jones, C., and R. D. Suttkus. 1972. Notes on netting bats 
for eleven years in western New Mexico. Southwestern 
Naturalist 16:261-266. 

36. Jones, C., and J. L. Paradiso. 1972. Mammals imported 
into the United States in 1969. Bureau of Sport Fisheries 
and Wildlife: Spec Sci. Rep. - Wildl. No. 147. ii + 33 pp. 

35. Jones, C. 1971. Notes on the anomalurids of Rio Muni 
and adjacent areas. Journal of Mammalogy 52:568-572. 

34. Jones, C., and H. Setzer. 1971. The design of a holotype 
of the West African pygmy squirrel, Myosciurus pumilio 
(Leconte, 1857) (Mammalia: Rodentia). Proceedings of 
the Biological Society of Washington 84:59-64. 

33. Jones, C. 1971. Wing loading in Plecotus rafinesquii. 
Journal of Mammalogy 52:458-460. 

32. Jones, C. 1971. Notes on hairy frogs ( Trichobatrachus 
robustus Boulenger) collected in Rio Muni, West Africa. 
Herpetologica 27:51-54. 

31. Jones, C. 1971. The bats of Rio Muni, West Africa. 
Journal of Mammalogy 52:121-140. 

30. Jones, C., and J. Sabater Pi. 1971. Comparative ecol¬ 
ogy of Gorilla gorilla (Savage and Wyman) and Pan 
troglodytes (Blumenback) in Rio Muni, West Africa. 
Bibliotheca Primatologica No. 13. iv + 96 pp. 

29. Jones, C. 1970. Mammals imported into the United 
States in 1968. Bureau Sport Fish, and Wildlife, Special 
Scientific Report - Wildlife No. 137. 30 pp. 

28. Jones, C., and H W. Setzer. 1970. Comments on Myo¬ 
sciurus pumilio. Journal of Mammalogy 51:813-814. 

27. Jones, J. K., and C. Jones. 1970. Dates of publication 
of numbers in the North American Fauna series (Letter 
to the Editor). Journal of Mammalogy 51:845. 

26. Jones, C. 1970. Stomach contents and gastro-intestinal 
relationships of monkeys collected in Rio Muni, West 
Africa. Mammalia 34:107-117. 

25. Jones, C.,T.W. Martin, andW. A. Mason. 1970. Survival 
of an escaped Callicebus moloch in southern Louisiana. 
Laboratory Primate News 9:6-7. 

24. Jones, C. 1969. Notes on ecological relationships of 
four species of lorisids in Rio Muni, West Africa. Folia 
Primatologica 11:255-267. 

23. Jones, C., and J. Sabater Pi. 1969. Sticks used by chim¬ 
panzees in Rio Muni, West Africa. Nature 223:100-101. 

22. Lowe, C. H., C. J. Jones, and J. W. Wright. 1968. Anew 
plethodontid salamander from Sonora, Mexico. Contribu¬ 
tions in Science, Los Angeles County Museum 140:1-11. 


12 


Clyde Jones Memorial Volume 


21. Jones, C., and J. Sabater Pi. 1968. Comparative ecology 
of Ceroooebus albigena (Gray) and Ceroooebus torquatus 
(Kerr) in Rio Muni, West Africa. Folia Primatologica 
9:99-113. 

20. Jones, C., and J. Pagels. 1968. Notes on a population of 
Pipistrellus subflavus in southern Louisiana. Journal of 
Mammalogy 49:134-139. 

19. Sabater Pi, J., and C. Jones. 1967. Notes on the distri¬ 
bution and ecology of the higher primates of Rio Muni, 
West Africa. Tulane Studies in Zoology 14:101-109. 

18. Findley, J. S., and C. Jones. 1967. Taxonomic relation¬ 
ships of bats of the species Myotisfortidens, M. lucifugus, 
and M. occultus. Journal of Mammalogy 48:429-444. 

17. Arata,A. A., andC. Jones. 1967. Homeothermy in Car- 
ollia (Phyllostomatidae: Chiroptera) and the adaptation 
of poikilothermy in insectivorous northern bats. Lozania 
14:1-10. 

16. Jones, C. 1967. Growth, development and wing loading 
in the evening bat, Nycticeius humeralis, (Rafmesque). 
Journal of Mammalogy 48:1-19. 

15. Jones, C. 1966. Changes in populations of some western 
bats. American Midland Naturalist 76:522-528. 

14. Findley, J. S., and C. Jones. 1965. Comments on spotted 
bats. Journal of Mammalogy 46:679-680. 

13. Jones, C. 1965. Ecological distribution and activity 
periods of bats of the Mogollon Mountains area of New 
Mexico and adjacent Arizona. Tulane Studies in Zoology 
12:93-100. 

12. Findley, J. S., and C. Jones. 1965. Northernmost records 
of some Neotropical bat genera. Journal of Mammalogy 
46:330-331. 


11. Findley, J. S., and C. Jones. 1964. Seasonal distribution 

of the hoary bat. Journal of Mammalogy 45:461—470. 

10. Jones, C., and J. S. Findley. 1963. Another record of the 
lyre snake, Trimorphodon vilkinsoni, in New Mexico. 
Southwestern Naturalist 8:175-177. 

9. Jones, C., and J. S. Findley. 1963. The long-nosed bat 
in New Mexico. Southwestern Naturalist 8:174-175. 

8. Findley, J. S., and C. J. Jones. 1963. Geographic varia¬ 
tion in the least cotton rat in New Mexico. Journal of 
Mammalogy 44:307-315. 

7. Findley, J. S., and C. J. Jones. 1962. Distribution and 
variation of voles of the genus Microtus in New Mexico 
and adjacent areas. Journal of Mammalogy 43:154-166. 

6. Jones, C. J. 1961. Additional records of bats in New 
Mexico. Journal of Mammalogy 42:538-539. 

5. Jones, C. J. 1961. Additional records of shrews in New 
Mexico. Journal of Mammalogy 42:399. 

4. Findley, J. S., and C. J. Jones. 1961. New United States 
record of the Mexican big-eared bat. Journal of Mam¬ 
malogy 42:97. 

3. Findley, J. S., and C. J. Jones. 1960. Geographic variation 
in the yellow-nosed cotton rat. Journal of Mammalogy 
41:462^469. 

2. Jones, C. J., E. D. Fleharty, and A. H. Harris. 1960. 
Unusual habitats of grasshopper mice in New Mexico. 
Journal of Mammalogy 41:275-276. 

1. Fleharty, E. D., and C. J. Jones. 1960. Possible Himalayan 
color pattern in Eutamia. Joumal of Mammalogy 41:125. 


Prologue 


“We learned about field work, and for some reason we loved it”: Oral History 
Excerpts from the Reminiscences of Clyde Jones 

David Marshall 


Part I: Recollections of Formative Years and Early Career 

Interview of 6 November 2003 at Texas Tech University, Lubbock, Texas 

Clyde Jones (CJ): I was born in Scottsbluff, Nebraska, March 3, 1935, a typical Depression baby... One of 
the remarkable things about my family—I was raised by older people. My mother was forty-two when I was 
bom, and at that time that was unusual. And I had an older brother who was nine years older than me, so I was 
obviously sort of a tag-along... 

Shortly thereafter my mother and father were divorced. My father remarried, and my mother never did. She had 
been a career school teacher in Nebraska, and she decided for some reason not to re-enter that profession. So she 
went into the cattle business with my unmarried uncle and my unmarried aunt and they raised purebred Hereford 
cattle. And we lived on a small ranch ten miles north of Burwell, Nebraska. We lived in the Sand Hills, and I had 
a great childhood. I thought and thought and thought and I can’t remember anything bad about my childhood, 
and I think part of it was that I was raised by older people. There were no other children in the family. I had 
two additional aunts who were school teachers and they married very late... they never had children. My uncle 
Bob, who became my surrogate father, he never married and my aunt never married. And so I was protected and 
babied by those people... 

I started school in a country school which was about three miles from the ranch headquarters. My brother and I 
went to school there, starting in the first grade, and I was sort of bored by all of that because I knew how to read 
and write and everything because of my school teacher aunts. On those long winter evenings they would read 
to me, and I would read to them... 

My brother dropped out of school and joined the Marine Corps, and he was killed in the battle of Iwo Jima. That 
had an impact [spoken emotionally]... 

Between the second and the eighth grade I skipped a year—I skipped a half a year and then another half a year. 
And so I was pretty young when I entered high school. I graduated when I was sixteen, and then my mother in¬ 
sisted that I lay out a year before entering college so I would again be with people my own age. And so I worked 
for the Navy at a defense plant building hundred-and-five-millimeter shells... 

Because I lived on the ranch, I was interested in prairie dogs, and gopher snakes, and things like that. I had a big 
gopher snake and I put him in a cage in my bedroom upstairs in the old ranch house and one day went upstairs 
and he was gone. And I sort of looked around, and my mother had some ladies down stairs in the living room 
and then my mother called. This plaintive voice, said, “Clyde, could you come down here please?” And what 
happened was my mother would have some ladies come visit her and then toward the end of the visit my mother 
would play the old pump organ that we had and they would sing, or she would just play. Or my Uncle Bob, Aunt 
Mary, and I would be summoned to come and sing and that was a real trip, you know. But there was this gopher 


13 


14 


Clyde Jones Memorial Volume 


snake lying on the keyboard of the pump organ, and so all of them were aghast at all of this, but I got him and put 
him back in his cage. That was the beginning of the banning of pets in the house. No more! Okay? 

David Marshall (DM): Did you hunt in the Sand Hills? 

CJ: Yeah, there was quite a bit of stuff, you know, a lot of water fowl, a lot of mammals on the ranch. And I 
had another—the uncle who married one of my school teacher aunts was an amateur taxidermist. And he knew 
most of the animals, and he collected. We picked up things, and he prepared them in his amateur way. They 
were mostly awful but we didn’t know that at the time... 

We had pocket gophers, they were common in the Sand Hills, and we grew grass. We grew purebred Hereford 
cattle and grass, and we had those old bar mowing machines, single-bar mowing machines. And the gopher 
mounds, the sand would wear out the plates on them, and so I was hired to go trap pocket gophers. My uncle 
bought me a couple of sets of Victor Gopher Traps and on the weekends I would ride around and trap gophers. 
And I didn’t know it but I was a mammologist. 

DM: Did you ever skin them or cut them open to see how they ticked? 

CJ: No, no, no, we just trapped them and poked them back down the hole and covered the hole, and buried and 
flattened out the mound, you know. 

DM: You did mention... that you butchered cattle on the ranch. 

CJ: Yeah, we butchered cattle and we butchered one pig. Let me tell you the pig story. My brother and I were 
out on our horses and we encountered this strange thing, and the horses were jumping around because they had 
never seen such a thing, and it was a pig. It was a red pig... and it had a paint bucket stuck on its face, and it 
was roaming around on our property. And we finally got a rope on it and we dragged it home and my uncle 
Bob came out of the bam and he kicked the paint bucket off of this pig. And I guess I was the first thing the pig 
saw—the pig imprinted on me and it just followed me everywhere and tried to follow me into the house and it 
just followed me everywhere. Finally, my mother let it come on to the porch but, you know, no pets in the house. 
That was a firm rule. Well anyway, I babied the damn pig. The pig grew up and my mother got on the phone 
and tried to find the owner of the pig and no one ever claimed the pig. So it grew up and we butchered it and we 
ate it. (laughs) But yeah, we butchered our own cattle, we had a big garden. Times were tough. We didn’t have 
much cash. We had the land, we had the livestock, we had the equipment, which we were constantly working 
with, but cash was a little—we had to sell something to get money, that’s simply how it worked. We had to sell 
something to get cash, and I guess that was typical of that era following the Great Depression. I’d like to say 
here that my parents—they never got over the Great Depression, they were sure it would come again, and maybe 
it has. They were sure that the Dust Bowl would come again. They were sure that World War I and World War 
II would happen again. These things always came up in sort of idle conversation around the dinner table, or on 
some Sunday gathering or something. Those things were never forgotten. 

DM: Was that area pretty well devastated during the Dust Bowl days? 

CJ: Yeah, I remember going out with my uncle and he would burn Russian Thistles [Tumbleweeds] that piled 
up in great masses against the fences. And we went to great expense to go to metal fence posts so we could burn 
the Russian Thistles that piled up against them. There were a lot of them. I remember that as a vivid experience, 
that in the Sand Hills, Russian Thistles sprouted and grew quite well. 

DM: Do you recall any wildlife observations? 


Marshall—Prologue 


15 


CJ: Deer were very scarce. Deer and wild turkeys were very scarce when I was a young boy. Now they’re very 
common in that area. Jackrabbits were common, bunnies were common, pheasants were common, grouse were 
still present. The best place to hunt grouse was, they would get on top of the haystack in the winter time, and so 
you just go out and shoot them, shoot all of them. And we would hunt a lot of waterfowl. The plan there was to 
put a gunny sack over you and crawl up on a playa lake and shoot, and you quit when you filled the gunny sack. 
We were supposed to be conservationists but we weren’t. But one of my uncles became appointed as a game 
warden. Well, that changed all the rules, (laughs) That changed all the rules immediately. He came and lectured 
to us and we learned about limits and things like that, and that changed everything. 

Bill Tydeman (BT): Well you mentioned... some favorite books that you had in your early years that you still 
have in your family library? 

CJ: Yeah I still have those. 

BT: And one is [James Gilchrist] Lawson’s WildAnimals: Photographs and Descriptions of 100 Important Wild Animals. 

CJ: Oh, it’s a wonderful book from that era. I still have all these at home. Yeah, it’s just a book with portfolio 
photographs of animals, important animals... 

I don’t know why I kept those things either, except those were the things I learned. I learned about mammals 
from those books, and those were all very non-technical kinds of productions. But it was great, for a kid like 
me it was great, okay? This was an inroad into knowledge about mammals that I couldn’t get, that didn’t exist 
anywhere else. There wasn’t a Mammals of Nebraska, or there wasn’t Mammals of the Sand Hills. There weren’t 
any of those things. Those things didn’t come along, my God, until the late Knox Jones wrote The Mammals of 
Nebraska in the 1950s. 

BT: The other thing that interests me, Clyde, in talking about your days at home and the influence of your mother, 
you mentioned that you had in a sense a family story hour, or a time in which each week you would tell stories, 
or talk about your readings? 

CJ: Yeah, she was very strict about that. She included herself—she and my two school teacher aunts who lived 
with us or they stayed with us on weekends. They lived with a family near their school, a country school where 
they taught during the week, but they lived with us on weekends. And we had reading assignments, to read some 
book and then after dinner we would report on it. And she was very strict about that, and I got into these mam¬ 
mal books and so that’s what I would read with a couple of exceptions: the Trail of the Loop, I think I mentioned 
there, and then along came a book titled Old Jules, which was a real classic about an early settler who came alone 
to the Sand Hills and it’s the story of his life. He had what, three or four wives? 

BT: Yes, and a daughter that becomes pretty significant— 

CJ: Yeah, his daughter Mari Sandoz became a very important writer in Nebraska history. Yeah, we had to report 
on these. I mean, and you didn’t miss a report, okay? That was another thing that didn’t—you didn’t say, “Well 
I don’t have anything to report.” You didn’t. This was one of those, as my uncle Bob called them, this was a 
non-negotiable. 

DM: I have the feeling that this reading was more inspiring than high school biology, from some of the comments 
you’ve made. Could you tell me a little bit about high school biology? Or what was the real impact on your life 
that would cause you to become a mammologist? 


16 


Clyde Jones Memorial Volume 


CJ: Well, let’s touch on high school very briefly. I learned one important thing in high school: I learned to type 
because my mother made me take typing. And as one of two guys in the class of about thirty females, you can 
imagine the names we were called. But here we are, two football players taking typing, and those old unmarked 
keys, a-s-d-f-j-k-l-semicolon. Jesus! How could you ever forget that? And that’s why I think, yeah, I think I’ve 
been fairly effective because I learned to type like a typist, rather than hunt and peck... 

I was sort of an undisciplined brat in high school. High school biology didn’t thrill me very much because I knew 
something about all of that stuff. I knew about reproduction, I knew about reproduction systems by watching the 
cows and the dogs. We had a bunch of dogs—you know. I knew all that stuff. I enrolled in college, and I had 
saved enough money to pay for the first full year of my tuition and my lodging. I lived in the dorm. 

DM: This was at Hastings? 

CJ: At Hastings College which is a Presbyterian college. And I went to talk to the football coach and he checked 
me over. He weighed me. I weighed one-hundred and eighty pounds, and he said, “What are you?” And I said, 
“Well, I’m a center or a guard.” He kind of laughed and said, “Yeah, he’s a center or a guard.” He said, “Here’s 
the line coach why don’t you talk to him.” And the line coach said, “Yeah, you’re a center to guard, you know.” 
He said, “Why don’t you snap the ball to me a few times.” And I did, and he said, “Yeah, you’re a center.” But 
he said, “You’re kind of small.” And I said, “Well, you know, I was an honorary mention all-conference in high 
school.” And he said, “I don’t care about that shit. I want to know what you can do in college.” And so we prac¬ 
ticed and the B-Team, which I was on, we had a game with an outfit at Norfolk, which was a junior college at the 
time. And we went there and he said, “Now, you guys are going to learn the difference between high school and 
college. This is your first game.” And the Korean vets [veterans] were returning. H e said, “There’re going to be 
some older guys and they’re going to be tough on you.” And we got in this game and the guy across from me, 
he just flattened me two or three times, and bloodied my face—oh, at that time we didn’t have face masks, okay? 
We just had interesting pots and no face guards, and I got my face bloodied, and the coach said, “Well, we know 
who’s looking around in there, you’re looking around in there, you’re getting your face bloodied.” And that guy 
was just flattening me, and I said, “Well, if you put me back in, I think I can take him.” And he just flattened me 
again. And they called a timeout and we were running to the sideline and this guy was just standing there, and 
I just gave him one as I went by, I hit him right in the neck and he just dropped, and it was a totally illegal and 
unconscionable thing but nobody saw it except my coach and he said, “You know, Jones,” he said, “You have 
possibility.” (laughs)... 

I played three years at Hastings. I lettered three years. I didn’t play my senior year because I discovered science. 
I used to go down to... a little museum in Hastings called “The House of Yesterday,” and it’s a typical mid-western 
city museum. It has farm equipment and all kinds of things. But I used to go down there, and I went down there 
so often they let me go behind the little exhibits and things. And there was a guy down there skinning birds, and 
I was fascinated by that. And I found out that you could make a living doing that. You could get paid to skin 
birds and skin mammals and things. And I went back to the college and talked to my advisor, who was a guy 
named Dr. Moulton—“Moldy” Moulton we called him. And he said, “Yeah, there is a field there.” But he said, 
“You have to take the following courses.” I had been going to be a history major, and I thought my future was in 
teaching history and coaching in high school some place, because that’s what you did in that generation. Women 
were either secretaries or school teachers; men were either ranchers or taught school and coached. I mean, those 
were the opportunities in that time, in that place. And so, in my senior year, he enrolled me for about twenty 
hours of biological sciences—comparative anatomy, all that good stuff. And God that was great stuff. I thought 
it was great. And Gene [Eugene] Fleharty and I had become friends in 1953, and Gene was a year ahead of me, 
and he went off to graduate school. He went to [University of] New Mexico. And he and Jim Findley found each 
other. And Gene influenced me to apply at New Mexico and my advisor John Moulten said, “Yeah, you should 
go to New Mexico.” He said, “They never do anything practical down there.” And I thought, well, that was an 


Marshall—Prologue 


17 


interesting thing for him to say. And I didn’t know about wildlife biology even though my uncle had become a 
game warden. I didn’t know what you did—I mean he was just a rancher and he was appointed a game warden. 
I thought, that’s the luck of the draw, I thought, that’s how that happens. But, Fleharty started filling me in on 
what he did and I talked it over with my mother and she thought I should go. She thought I should go to New 
Mexico because I could learn about different cultures. I could get out of the Sand Hills, and besides that she 
wanted to come visit me, and so, okay. 

DM: What about Fleharty, was he from right around Hastings? 

CJ: He’s from Hastings. He comes from a family of six boys in that family. 

BT: Was he on a clear biology track as an undergraduate? 

CJ: No, I don’t think he knew what he wanted to do either. But John Moulton encouraged us to apply to graduate 
schools, and so he was accepted at New Mexico and he went there. And I was accepted at, let’s see, some place 
in Illinois but I didn’t really want to go there so— 

DM: Didn’t you teach high school for a little brief period too? 

CJ: As a practice and a substitute teacher. 

DM: While you were attending Hastings? 

CJ: Yes. And I quickly decided that high school teaching was not for me. I quickly decided that’s not my bag... 

So we went to New Mexico, and Jim Findley loves to tell this story: I walked into his office wearing a shirt with 
the sleeves cut out and an old cowboy hat on and a pair of boots. And Findley tells the story that I still had horse 
shit on my boots. I don’t think that’s true. And he said, “What do you want?” And I said, “I came to get my 
PhD.” And Fleharty was with me, and Findley looked at my transcript from Hastings College... 

Findley took a look at my college transcript and he said, “You know if you were here you would hardly be a senior.” 
I didn’t take chemistry in college... I never had botany, either, as an undergraduate because botany wasn’t taught 
at Hastings. They didn’t have one. They had a one-man biology department, one-man biology and geography 
department, and so, we were a little weak in background information. 

DM: But, you know, they produced two biologists. Is that just coincidence? Are there others that went into 
biology at Hastings? 

CJ: They produced others—there’s a guy visiting us from Lincoln, Nebraska, named Hugh Genoways. He’s a 
graduate of Hastings. They have produced half-a-dozen mammologists. 

DM: Why is that? 

CJ: We don’t know. We don’t know. And I guess the track that Fleharty and I led sort of attracted a couple of 
other people. It attracted Hugh Genoways, who [thought] “I’ll follow in the footsteps of those guys.” And he did. 
But as Findley put it, “I had to make something out of two jocks.” (laughs) And I don’t know if he did or not 
but we survived. I think my first semester as a graduate student I think I got four hours of graduate credit, all the 
rest were leveling courses—botany, chemistry, stuff like that. Jesus! I got a C in one of my chemistry courses, 


18 


Clyde Jones Memorial Volume 


and the rules were: if you got six hours of C you were out. And Findley said, “Well, you got that out of the way; 
now you don’t have to get another C.” That was his positive approach to it, you know, (laughs) For God’s sake! 
Actually, I was very happy with my C at the time. And we learned about trapping and skinning mice. And we 
learned about field work, and for some reason we loved it... 

We did a lot of field work. NSF [National Science Foundation] came out with a program of grants for terminal [de¬ 
gree] people to finish their research and we both applied for that, and for Christ’s sakes, we got them. And they paid a 
bunch of summer money to us, so we gave the money to our wives and we went and lived in the field. Gene lived at 
a place called Wall Lake. He was studying garter snakes for his PhD. I lived at a place called Willow Creek because 
I was studying bats. Bat nets had just become available. And we didn’t really know what we were doing, okay? 

DM: Before bat nets became available was there anyone doing that kind of work? Were they finding a way to— 

CJ: The only way to get bats was to shoot them. And out of a case of shells you might get two or three, because 
they’re pretty quick, you know. The other way to get them was to stretch a wire about that far above the surface 
of the water and they would come in drinking and fly into it, and if you were quick enough, you could get them. 

DM: Did the use of bat nets really promote the study of bats? Did it have a real significant impact? 

CJ: Yeah. The study of bats exploded; it exploded with the availability of bat nets. 

DM: And you were right there? 

CJ: Yeah, we were on top of it. And science was funny then too. Remember, this was—statistics hadn’t been 
invented yet, computers hadn’t been invented yet, things were primitive, and the whole concept of hypotheses 
hadn’t been invented yet. The approach was, we go into the field with a sack and a stick and collect everything 
we can collect, and then bring it back and try to decide what to do with it. It was that approach, and it worked. 
We were working with Jim Findley on the Mammals of New Mexico. And so we were collecting, and we were 
collecting everything in sight, literally. We were shooting it or trapping it or netting it. We were collecting 
everything. And I became interested in bats because I stopped in at the University of Arizona, and E. Lendell 
Cockrum—who’s still alive, incidentally—he was a bat person there, and he said, “You ought to study bats.” 
He says, “There’s a real need for it.” And I thought, yeah, I ought to study bats. And I convinced Findley that I 
should, and he was a little nervous about that at first. And it turned out that I lucked into something. I caught a 
whole bunch of bats and it worked out that different species forage at different times and, therefore, they feed on 
different things. And that was sort of the beginning of resource partitioning as we now know it. And we didn’t 
even call it that then. We just said, “Yeah, these bats feed at this time, these bats feed at this time, and these bats 
drink at this time, and these bats drink at that time.” You know the phrase “resource partitioning” hadn’t been 
invented yet either, (laughs) 

DM: Where were you doing most of your bat netting at this time? 

CJ: At various places in the mountains of west-central New Mexico. I lived in a tent at Willow Creek and trav¬ 
eled around to several places and netted bats... It’s in the Mogollons. And Fleharty lived at Wall Lake. And about 
every two weeks I would go see him or he would come see me. And the road from Willow Creek to Wall Lake 
was a real bitch, and if it rained, it was worse than that, it was just solid mud. 


DM: Can you describe your camp life? 


Marshall—Prologue 


19 


CJ: I don’t think we were ever lonely, maybe a little bit sometimes, but we were busy. First of all, subsistence 
was important. We were busy feeding ourselves and keeping ourselves dry. And in the mountains it would rain 
frequently. In the summertime, the summer monsoons came through, and August was rainy. It would rain. And 
he was working on garter snakes and he was in the lake a lot. I was netting bats a lot. We didn’t have waders at 
the beginning. Later on, I got a pair of chest waders and they were wonderful. But yeah, we were just busy. We 
would—I would net bats all night and then skin bats all day. 

DM: What about food and shelter? 

CJ: Well I lived in a tent. I lived in a ten-by-ten umbrella tent, and I’d cook a good meal maybe once a day; lunch 
the rest of the time. Peanut butter sandwiches are really good, you know, they’ll carry you a long way. (laughs) 
Yeah, you can live a long time on peanut butter sandwiches. Peanut butter is also rat bait. 

DM: When you launched into this field work, early on, did you feel that you had found your niche, or did you 
question whether this was really what you wanted to do? 

CJ: This is what I wanted to do the rest of my life, was trap and bag mice, and net and bag bats. This was it. 
And I discussed this with my mother. And as I comment in there, I’m not sure she ever really understood what 
and why I did what I did, but she supported it... 

But yeah, this is what I wanted to do, and I wanted to teach other people how to do the same thing. I wanted to 
teach and do this. And then I finished—well, let me tell you another story. Fleharty and I were roaming around the 
mountains, and I was driving. Jim Findley loaned us his old Chevrolet pick-up, and he had a camper on the back 
made out of plywood, you know how those are... It was an awful beast. We took a trip and went into the Black 
Range of New Mexico and we patched the tire with hand tools, and patched the tire with a sleeping bag patching 
kit—and the damn thing held! (laughs) And so we went on with our trip and I was driving... and he says, “Stop, 
stop, I just saw a white chipmunk.” Yeah, yeah, yeah, white chipmunk? White elephant yes, white chipmunk, 
no way. He says, “Back up, back up, I saw, I’m telling you, I saw a white chipmunk.” And I thought “This guy, 
he’s nuts.” He says, “Back up goddamn it, I saw a white chipmunk.” So I backed up, and sure enough, there was 
a white chipmunk sitting on a rock and he shot it. And we skinned it and stuffed it. You know, whoever heard 
of a white chipmunk? So we got back to Albuquerque and—oh, we got to Silver City and a guy at a Standard 
filling station there sold us a tire on the promise that we’d pay him the next time we came by. How about that? 
We had like five dollars and he said, “No, I’ll just sell you a tire and next time you come through, I recognize the 
truck, the next time you come through you pay me for the tire.” It was like thirty dollars or something. And we 
thought, “Hey, this is great, we should have bought all four of them.” (laughs) And we called Findley, called him 
on the phone from this service station, and he said, “Now what have you done?” And we said, “Well, we bought 
a tire, and we will have to pay for it the next time we come through Silver City.” “Yeah, yeah, yeah, give me 
another bullshit story.” “Well, we shot a white chipmunk.” “Great guys [Findley said in disbelief], when are you 
coming home?” “Well, we’re starting home right now, we will get there tomorrow.” And so we wrote a paper 
on this white chipmunk and submitted it to the Journal of Mammalogy, and it was accepted. And we thought, 
“Hey, Jesus, this is really neat, you know, you can publish on what you do.” And so we went to Findley and we 
said, “Well, there’s this thing about buying reprints, and we don’t have any money.” And he said, “You go to the 
chairman and you ask him to buy your reprints.” And he set us up. So we walked in—we made an appointment, 
we walked into the chairman’s office and we said, “We wrote this paper on this white chipmunk, and we have 
to buy some reprints, and we wondered if the department would buy reprints.” And the chairman just looked up 
at us and he said, “Graduate students don’t publish.” And we thought, “What the hell was this?” And Findley 
was waiting right outside the door laughing his ass off. He said, “I’ll buy the reprints; I just wanted you guys to 
get the departmental philosophy.” That was the philosophy: graduate students don’t publish, (laughs) And that 


20 


Clyde Jones Memorial Volume 


was the beginning of it all—hey you can publish this stuff, and you can publish it in the Journal of Mammalogy, 
shit, we’ll be famous, (laughs) That’s what we said to each other, “We’ll be famous.” And Findley said, “Yeah, 
yeah, you guys are already—‘infamous’—is the word.” Geez, but that was the philosophy. Graduate students 
don’t publish. So— 

BT: But at the time that you’re at New Mexico doing your graduate work, and you’re talking about the methods 
and practices that existed then, I mean Watson and Crick are just coming out, the double helix, and the— 

CJ: Oh God, did I luck out on that one. Yeah, I was taking my final exams for my PhD.... The night before 
my oral exam, I went to the drug store to buy a relaxant. And there while waiting to checkout, there was a Life 
magazine with the double helix on it, and a big article about Watson and Crick. And I bought the magazine. I 
thought, “I better buy that.” And I read that damn thing that evening and the next day at my oral exam, the first 
question out of the box, Jim Findley said, “Have you ever heard of DNA?” And I thought, “I’ll just get the bas¬ 
tard.” I jumped up and I drew the double helix on the blackboard. I talked about it for about three minutes, and 
Findley said, “You son of a bitch, you found the magazine.” (laughs) That was in the early sixties, Jesus. It was 
luck, just luck. Just blind luck all of my life... 

DM: Was it Findley that got you the museum curator position at UNM? Didn’t you work there in the museum 
for a while? 

CJ: Yeah, Findley got that job for me. That was great... I finished my PhD and I was hired as the assistant cura¬ 
tor. I was the assistant curator for mammals, birds, fossils, plants, reptiles, and amphibians. I was busy. 

DM: What kind of work did you prefer: the teaching, or the museum work, or the field work? 

CJ: Well, field work of course number one; it still is. Museum work is very important. Teaching is a way to 
achieve those things... 

My mother was very, very supportive of me going to graduate school and becoming a college teacher. She was 
very supportive of that, in every way. I can’t think of a single incident that she was not supportive of. She was 
extremely—almost pushy—supportive that I would find myself, I would be a college teacher, and I would have a 
better life than she had. I heard that many times. So, I had the assistant curatorship at Albuquerque and I realized 
that I would just always be Clyde the graduate student, that I had to get away from there, I had to leave, I had to 
go someplace. And I applied around and interviewed at two or three places... 

Fleharty had left. He went to Nebraska Wesleyan in Lincoln, Nebraska, for one year. He was paid five thousand 
dollars and we thought, “Hey, hey, we’re on the right road here.” I got a temporary job at Tulane University to 
replace Norman Negus, who was on sabbatical for a year, and I was paid eight thousand dollars. And, Jesus, 
everybody was hanging around me, you know. And so we moved to New Orleans, moved to New Orleans in my 
Dodge with a small U-Haul trailer with everything we owned, and lived in a little apartment on campus; finally 
found an apartment of our own out by Lake Pontchartrain, and lived out there. [I] taught eighteen contact hours 
and [would] come dragging into my office at nine o’ clock in the evening. And I picked up a couple of graduate 
students, picked up three graduate students in fact, and they would say, “Why do you come to your office at nine 
o’ clock?” And I said, “Stupid, that’s when my last class ends.” I taught my ass off at Tulane. 

DM: What kind of an adjustment was that, having grown up in Nebraska, lived in Albuquerque, and then moving 
to New Orleans, of all places? 


Marshall—Prologue 


21 


CJ: That was an experience. Albuquerque was a very cosmopolitan city and the University of New Mexico was 
a very interesting place. All the news media focused on Berkeley [University of California], Berkeley didn’t 
have it, man—we had it, but we were a small place, okay? I mean, we had everything, mixed marriages, mixed 
couples, this was the hippie era. We all grew hair. We were going to strike once for—oh, because of the dress 
code. And we pushed this all the way to meet with the President of the University of New Mexico. And we had 
a spokesperson and everything, and he met with us, he was a guy named Tom Popejoy. And he met with us, and 
he was very patient. He listened to us and he said, “You know, I’m going to take this under advisement; I’m going 
to think about it for a month. I’ll meet with you a month from today.” And he said, “Oh by the way, I’m going 
to freeze salaries until a month from today.” And he left, and we looked at each other and we said, “This isn’t 
working out.” (laughs) “This is not what we had in mind.” And we met with him a month later, everybody had 
clothes, everybody had a shirt with a collar, and everybody had socks and shoes on. Sometimes one blue sock 
and one grey sock but everybody had clothes. And he said, “You guys are a good looking bunch of guys, you are 
going to be paid, here are your checks.” (laughs) Jesus! That was our learning experience about organizations 
and striking and unions. And that lasted with us, we have never belonged. Fleharty and I have never belonged 
to a union or an organization like that since. We learned—he was great, he was a great President... 

When I arrived at Tulane, Tulane had integrated itself racially and sexually, and that was interesting. And living 
in New Orleans was very interesting also for a guy from the Sand Hills... 

I became acquainted with the director of the Primate Center who was a guy named Art Riopelle. And I don’t 
know, for some reason he liked me too, and he said, “You ought to come to some of our seminars.” And so, okay, 
I’ll go to some of his seminars, you know, I don’t have enough to do already. So, I went to one and it was a girl 
[speaker] named Jane Goodall. And he introduced me to her and her husband who then was Baron [Hugo] van 
Lawick. And we talked about her work in Africa, she gave a seminar on her chimp work, and I was just totally 
enthralled with this. And she said that the late Mr. [Louis] Leakey was looking for somebody to study primates 
in West Africa—lowland gorillas. And I didn’t think anything about it. We talked about Dian Fossey and her 
work with highland gorillas, and I didn’t think anything about it. And she went away, and I went back across the 
river, across Lake Pontchartrain... I was beavering around Tulane and Art Riopelle called me and said, “Why 
don’t you come over and see me? Why don’t you come over and visit with me?” Okay, so I went over and he 
said, “I think we ought to apply for a grant; I think we ought to apply for a grant to send you to West Africa.” 
And any young mammologist that doesn’t want to go to Africa is not worth anything, okay? I wanted to go to 
Africa, but I thought, “This is unreal, this can’t be happening.” And he said, “Yeah, I’ll draft the grant, you fill in 
the information about yourself, and we’ll get some money from the National Institute of Health, and we will get 
some money from [the] National Geographic [Society], and we’ll send you to Africa for a year.” And I thought 
“Okay, I’ll take a shot.” So we put the grant together, and we submitted it, and, quite frankly, I sort of forgot 
about it. And that summer [Royal] Suttkus taught—it was the summer of 1965. Suttkus had an environmental 
training grant, and he took a group of students from Florida to San Diego, and I went with him on that trip. God, 
it was great. We collected the shit out of everything. We collected a thousand mammals; a thousand specimens... 

And I got home, and my wife said, “You’re to call Art Riopelle immediately.” “Okay.” So I called him and he 
said, “Hey I have good news for you, we got the grant!”... 

So, in the summer of 1966,1 went to Africa. I went to Spain and met the Spanish coordinator who was the director 
of the Barcelona Zoo, and his biologist who was Jordi Sabater Pi, and I slogged around Spain. It was a wonder¬ 
ful place, wonderful beer in Spain, wonderful. San Miguel [beer] is just for your taste. You can get it here... 

And I went to Africa by myself. Jesus! Went to Rio Muni [present Equatorial Africa] and met the people and 
the army and everybody. It was a Spanish colony, and so I met everybody. Jesus Christ! What an operation and 
what a place to get into and to get out of. 


22 


Clyde Jones Memorial Volume 


BT: You mean all kinds of bureaucratic regulations and things that just—? 

CJ: Oh God—yeah, and there were essentially two governments in Rio Muni: the Spanish government, and 
they had what they called an “autonomous government.” They were training a government to run the place, 
because it was going to become independent. You know, you deal with the Guardia Civil, which was the army 
government that ran the place, and then you had to deal with the civil government, and that was a nightmare. 
Bureaucracy rampant, and double paperwork, and “Oh, that doesn’t work, you have to fill this out,” and so on 
and so on. Anyway, I slogged around Rio Muni and selected a couple of places I wanted to study. And I had 
to arrange to rent a house and ordered a car, ordered a Land Rover, which would be delivered sometime in the 
future, and incidental things like that. Checked out the medical facilities, caught malaria, had a great time. I 
went to get my—I’ll tell you a small story—I went to get my paperwork to leave and you had to have a blood 
test, a test for malaria, before you could leave the country. So I was standing in line, and I got up front and the 
guy taking the blood had a lancet, and he would take somebody’s blood and then he would wipe it off with an old 
rag. And I said, “I don’t want you to do that, I want you to use another lancet.” And he said, “This is the only 
one I have.” And I went to my Land Rover and got in my skinning kit and punctured my finger. I didn’t want 
hepatitis on top of everything. And I came back and taught in the fall semester at Tulane... And so I went, took 
my family to Africa in January of 1967... 

I had been there in the summer of ’66 and this was in January of ’67. Went to Spain, went to Africa from Madrid 
to Fernando Po, it’s now called Macias Nguema, the island on the west coast. That was a ten-hour prop plane 
flight, and everybody was sick, everybody barfed at least once on that flight. Flying over the West African desert 
was like that in a prop plane. Landed at Fernando Po, and I had convinced my son [Craig] that the reason to 
go to Africa was that he wouldn’t have so many Joneses around him. In New Orleans there were thousands of 
Joneses, okay, thousands of them. And so we walked into the terminal at Fernando Po, and there’s this tall black 
guy standing there and my son walks up to him and says, “My name’s Jones, how are you?” And the guy says 
[in a British accent ], “I say, my name’s Jones also.” (laughs) Craig says, “I want to go home.” (laughs) It was 
a British-educated Nigerian. [It was] really strange to hear that accent, incidentally, really strange. 

DM: How many total months did you spend in West Africa? 

CJ: Well, we spent a total of nineteen months... We were supposed to spend a year and a fortunate thing hap¬ 
pened. The Spanish devaluated the peseta. So, suddenly, I had more money than I thought I had, and I wired 
Riopelle and he contacted National Geographic, and they said, “Stay, take the money and stay.” On the way there 
I stopped in Washington and went to National Geographic where I met Dian Fossey for the first time, and they 
had a wonderful director of research [Leonard Carmichael]. He was the former secretary of the Smithsonian, and 
he said, “I’m not interested in glossy-paged articles, I’m interested in science.” And I thought, “This guy knows 
what he’s talking about.” And so, we were in Africa, the house I had rented, the furniture I had arranged for was 
still there, the brand new Land Rover was there—diesel Land Rover. If you have never driven a diesel Land 
Rover you haven’t experienced anything. It’s a diesel jeep, oh God, bump, bump, bump, bump. But Rio Muni is 
a small country about seventy-five miles wide and a hundred and twenty-five miles long and about five-hundred 
thousand people; about twenty white people when we were there. They thought at first we were French, or Ger¬ 
man, and when they found out we were American, we were very popular, because we were the only ones there. 
And they thought we were something special, I guess, I don’t know. They had all kinds of questions about life in 
America. And when I worked, I would go out and arrange to stay in a village, and hire a woman, stay in a nice 
house or some place, and hire a woman to boil water and to cook for me, and hire a boy to keep the goddamn goats 
off of me. (laughs) They had thousands of goats and they were everywhere. In my bed, in my car, on top of the 
Land Rover, they were everywhere. I had to hire some young man to keep the damn goats away from me. Jesus! 


Marshall—Prologue 


23 


DM: What were you trapping? 

CJ: Well, I was studying primates. 

DM: Just primates? 

CJ: And I wrote The Bats of Rio Muni and I wrote numerous—I wrote nineteen papers from the nineteen months 
I stayed there. That was my goal, and I achieved it... 

BT: You mean you went in with the idea of a paper a month? 

CJ: Yeah, I went there and said “I’ll write a paper a month.” And I achieved that goal, nineteen papers in nine¬ 
teen months. 

BT: Did you begin to think about, well maybe primates is what I want to do, to do primate research, or did you 
see it just as a separate path? 

CJ: I looked at primates from a zoologist point of view. And they’re really interesting, and there are some really 
interesting problems. But I came to the realization that those problems will never be solved, because primates 
remind you of little people and there’s a lot of emotion tied to getting permits to study primates. What really 
ought to be done is, somebody ought to go shoot about a hundred of them of every species, but you could never 
get a permit to do that. And the specimens that are in museums, the British Museum, the National Museum [of 
Natural History], the American Museum [of Natural History], Chicago [Field Museum], they are all caliper-worn 
from being measured for a hundred years, and they need new material to study the problems, and you would 
never get a permit to get it. 

DM: On a broader scale, do you have that problem with mammals where you wouldn’t have it with herps? 

CJ: Yeah, it’s a problem with mammals, it’s a problem... 

The African experience was a wonderful experience scientifically; it was a wonderful experience personally. The 
political problems were incredible. There was little to do in Bata for entertainment. Well, going to the market was 
entertaining and problematic, of course. They had a movie theater. The movies were all in Spanish. We went 
to the movies a lot. They had soccer games there; they played teams from Cameroon and Gabon and Nigeria. 
We used to go to soccer games—we were sort of special there because we were the only white people and my 
wife was the only woman there. It was a men’s thing, but we went. One time one of my trackers came by and 
he said, “Don’t go to the soccer game Sunday.” I thought, “Well, he wants my tickets or something.” He said, 
“No, please, don’t go to the soccer game Sunday.” Well, as it turned out, during intermission they brought out 
some political prisoners and shot them, and he knew that was going to happen. Everybody knew everything. I 
thought—I used to joke and say it must be drums, because something simple would happen—the license tag fell 
off my Land Rover, and I didn’t know it, and I heard about it for about a hundred miles. And I thought, “If I 
don’t recover that license plate, everybody is going to go crazy, they all know about it.” It’s a fantastic society. 
You get inland just a little bit and there are people there that have never seen the ocean, and the whole country is 
one hundred and twenty-five miles. And they’re very clan-oriented. They are called the Fang, it’s a sub-group of 
African culture. It’s very interesting. I liked them. I learned to like them and respect them, and yet I feel sorry 
for what they are subjected to and they are still subjected to incredible human rights problems. But it’s a small 
place, they don’t have anything. They discovered oil in 1995,1 believe, and oil companies pour some money 
into the country. Most of it goes to the dictator—it’s a mess. 


24 


Clyde Jones Memorial Volume 


DM: Did you happen to write anything about the society while you were writing about— 

CJ: I wrote a little bit about it, just a little bit. Yeah, my daughter says she thinks I should write a book about 
that, and I don’t know if I will or not. It’s a wonderful—the people are wonderful once you get to know them. 
They’re wonderful and they are clever and they can do wondrous things with their hands. I went down a river 
once with a dugout canoe with an outboard motor on it, and the motor quit, and we beached the boat. And this 
guy, with a Swiss Army knife, took the motor all apart, laid all the pieces out, and I thought, “Oh Christ, we’ll 
never get out of here, we’re in the mangrove swamps.” He laid all the pieces out on a rag and wiped every piece 
off with a rag, and put it back together and cranked it up, and I was amazed at this. I couldn’t do that; I couldn’t 
do that. They’re amazing. 

DM: Did you happen to keep journals during your field work? 

CJ: Yeah, I have field journals of all that stuff... That was a wonderful period of time for us and the children 
were very young, but they still speak Spanish; that’s all they had. We returned to the Primate Center and bought 
a house in Mandeville, Louisiana, and enrolled my son and daughter in the public school there. And one day I 
got a call at the Primate Center that the principal of the school wanted to talk to me. And I wandered in there 
and he said, “I have to talk to you about a problem. Your son and daughter are lining up with the black kids at 
the black drinking fountain.” He said, “Are you a blockbuster, or what are you trying to do here?” And I was 
absolutely flabbergasted by this—this was in 1968. And I did not realize that they still had a white and black 
drinking fountain, and I tried to explain to this guy that we spent nineteen months in Africa; the only children my 
kids knew were black children or Spanish children; they don’t know any better, they don’t get the segregation 
part, they don’t know any better, it’s purely an act of innocence. And I don’t think he ever really believed me. 
And I talked to Sherry and Craig about this and they said, “Well, I mean, we were talking to so and so, and she 
went to get a drink, and I went with her.” And I said, “That’s perfectly fine, I want you to do that. I just want 
you to know that things are a little different here than they were in Rio Muni.” And my son had sort of a wise 
comment. He said, “Yeah, things were better in Rio Muni, weren’t they Dad?” And I said, “Well, in some ways, 
yes, they were, yeah.” I couldn’t believe that. That was in 1968... 

I came back from Africa and was housed at the Primate Center and I had from National Geographic funds to 
write for a year. And I was looking for a job and I saw this ad with the Fish and Wildlife Service, and I applied 
for the job, and went for the interview and lucked out and I got the job. 

DM: This was Chief of Mammals— 

CJ: Chief of the mammal section of the old Burton Mammal Laboratories, which is a remnant of the Biological 
Survey that C. Hart Merriam operated. And, my God, I lucked out, I got this job. And it paid $13,300 for a year. 

DM: And that started in 1970? 

CJ: Yes. 

DM: How long were you in that position? 

CJ: Oh, I spent ten years there, but I became the director of the lab and I don’t remember exactly when that 
happened. And then, after ten years, the Fish and Wildlife Service moved me to Denver and combined the old 
Denver lab with the lab in Washington. And I lasted there until 1982. President Reagan appointed a gentleman 


Marshall—Prologue 


25 


named James Watt to be the Secretary of the Interior, and he came to visit me and I decided I should search 
elsewhere, (laughs)... 

I was very happy. I had a wonderful time in D.C., a wonderful ten years there. I had enough. And I’m a western 
person. I had enough of the government structure. Don’t misunderstand me; those were wonderful years for me. 
I made real lasting associations with a lot of people. I got to hire a lot of people. Our outfit grew and bloomed, 
especially under the Carter administration. He understood in part what we were trying to do, and yeah, I got to 
hire and see some wonderful people grow and develop, some of whom are still there. It was a wonderful ten 
years. I just wanted to move back to the Midwest... 

I loved it at Denver, okay? I loved that job. I had about two hundred people working for me. My management 
style is sort of different. I had an executive secretary, an assistant director, and a business manager; they reported 
tome. The assistant director took care of everybody else. It was a wonderful job. We opened and developed field 
stations in Haiti, the Sudan, Philippines, all over the world. In Alaska, we had field stations in Anchorage and one 
at Fairbanks; had the California situation. The Marine Mammal Act was passed. The Fish and Wildlife Service 
portion was given to me. Suddenly I had four million dollars and not a marine mammologist on the staff, and so I 
hired one. And he hired others and it was great fun, it was just great fun. When I left, we had two hundred people 
and about twenty-million dollars plus a whole bunch of money from AID [Agency for International Development] 
for the foreign field stations. I hated to leave Denver. The option was to move back to Washington. I and my 
then-wife were not terribly enthralled with that idea. I mean, we had spent ten years there; it was enough. We 
wanted to be in the west somewhere... 

And the late Knox Jones called me and he said, “I don’t know if you’re interested, but,” he said, “There’s a posi¬ 
tion coming open at Tech.” He said, “I have it that the museum director is going to resign.” And I thought, “Huh, 
I could do that.” Sol applied and I lucked out... 

I was interested in returning to work in the Chihuahuan Desert. And I came here as director of the museum. The 
so-called museum support groups were very strong because of weaknesses on the part of former directors. I put 
up with that for a while, and then I decided, “Bullshit, I’ll just go back to biology and be a professor and work 
on mammals of the Chihuahuan Desert,” which I did, with a fair level of success... 

Within my lifetime we’ve gone from no techniques, we’ve gone from where data overshadowed the technology, 
to now where the technology overshadows the data. This is a tired old phrase: “We need more data.” We went 
from having to hand-make distribution maps to making them with the touch of a key now, and there are a lot of 
holes, there are a lot of gaps, there are a lot of things we don’t know. There’s a collection of Chihuahuan Desert 
mammals here, we have, and other things. We have a hundred-thousand specimens. What do we know about the 
Chihuahuan Desert? Not much, not much—and it’s a very crucial area now, with the issue of water. 


Part II: Further Thoughts on Living and Working in Rio Muni 

Interview of 27 February 2012 at the home of Clyde Jones, Lubbock, Texas 

David Marshall (DM): Can you tell me the circumstances that took you to research in West Africa? 

Clyde Jones (CJ): Yeah, I was a young professor at Tulane, and I had finished my graduate work from New 
Mexico working on mammals of the southwestern desert, and I became acquainted with Arthur Riopelle. He was 
the director of the former Delta Primate Research Center, and he was a young man who was a psychologist, of 
all things. And he said, “Let’s do an African study.” And he had made a contact with a person at the Barcelona 


26 


Clyde Jones Memorial Volume 


Zoo and he said, “Yeah, we could get a Spanish contact and work in the Spanish colony in West Africa... ” And, 
well, I knew a lot about Africa. I knew where it was, geographically, and I had read “Tarzan,” and that’s about 
it. And a great surprise to me, we obtained funding. We applied to the National Geographic Society and to the 
National Institutes of Health, and both of them were approved... So, I went to Rio Muni, in the summer of 1966... 

He had also made a contact with Louis Leakey, who was a mentor of Jane Goodall and Dian Fossey, and he 
brought Jane Goodall to the Primate Center to give a talk... 

The primate center had a colony of captive chimps which Arthur Riopelle had studied, but he was interested in 
Lowland gorillas and chimps, and the ecological separation between the two. And so, I went to Rio Muni with 
the understanding that I could study other mammals as they came available to me. You know, I’d never seen a 
gorilla before. So I went to Rio Muni. Jordi Sabater Pi was there and he was familiar with the language. He 
spoke some Fang... One of the first things I learned was that most of the Fang that lived out in the villages were 
frightened of the rainforest. Their lives were centered around the village, and the slash-and-burn agriculture. 
And most of the men, women, and children never ventured into the rainforest. They had all kinds of mythical 
tales about what would happen to them if they went into the rainforest. Like they would be chased by a gorilla 
or chimp-like monsters, or be killed, raped, or pillaged. You know, all kinds of weird stories... 

But in most villages there was a hunter who went into the rainforest, and he captured bush meat. And he captured 
monkeys and small dik-diks, and all kinds of animals and birds. They ate everything. They ate fruit bats; they 
ate birds; they ate mammals; they ate everything. And it was usually a hunter, so I eventually learned to find one. 
It was usually someone who stood around the outside, and he would come around later and say, “I’m your man.” 

DM: So he wasn’t in society; he was kind of on the periphery. 

CJ: He was kind of on the outside. 

DM: Why was that? 

CJ: Well, because he went into the rainforest. And he was considered a brave, very brave man. And for some 
reason, they were armed with crossbows and blowguns and the ever present machete. And then all the hunters 
smoked a pipe. And I soon realized what it was, it was marijuana, which there was a patch at every village, and 
that was called an anti-fear device by me. These guys smoked the pipe and then went in to the forest, and I went 
with them on a hunt a couple of times. And they used the blowgun, and a poison dart, and they shot a monkey 
up high in the tree, and the monkey came crashing down. And he cut a slot in the arms and legs and put them 
together, and put them on his back, and put one of them on my back, and back we went to the village. And I 
was worried about the poison dart that might be somewhere. But what he did was, he felt all over, and found the 
poison dart, and extracted it. 

DM: Now what about the poison in the system, and then eating that monkey? 

CJ: Now that didn’t seem to bother them. But they had a way that they went at it, especially the pygmoids. 
When they got a monkey, they went at it by opening it up and eating the stomach contents. That was the first 
thing they did. 

DM: Raw? 


CJ: Yes, just cut it open and eat it, ghastly stuff. 


Marshall—Prologue 


27 


DM: You tasted this? 

CJ: Yes, it was terrible stuff. 

DM: This is what you said looked like split pea soup? 

CJ: Oh yeah, they ate it. They kind of danced around, and you know, scooped it up with their hands and ate 
it. They didn’t have—the people that lived in the woods— the pygmoids, or they were called the Bajeles —they 
didn’t have any utensils; they ate everything with their hands—everything. 

DM: Did they eat everything raw or did they cook some? 

CJ: They cooked sometimes; they cooked some meat or some things, but they just ate it with their hands. 

DM: And they ate it with gusto, the contents of the stomach? 

CJ: Yeah, oh yeah, it was a ceremonial kind of thing. 

DM: Now what was the ceremonial part of it? You said they got excited. 

CJ: Yeah they were excited, they danced around you know, and the guy would hand some to somebody, and he’d 
take it and eat it, and yeah, it was some kind of ceremony. I never did figure out their language, but for some 
reason that one little woman just sort of adopted me, and she took care of me, and she was quite observant. She 
observed me sitting on a stump or something, writing my field notes, and I went out with the guys and they came 
back, and there was a little table and a little stool, and she indicated that that was for me... 

DM: Were the pygmoids hospitable generally, or was this an individual characteristic? 

CJ: At first they were very standoffish, and, well, they’d never seen anything like me and they were quite stand¬ 
offish—but in time, they came around. She quickly adopted me. Well anyway, I sat on this stool, and of course 
I smashed it flat and my feet came up and broke the table all to pieces... because it was made out of little sticks, 
made for them, not for me. And they threw themselves on the ground laughing. They rolled on the ground laugh¬ 
ing, all of them laughed, and that was the funniest thing they’d ever seen. And I laughed too. When I laughed 
they laughed. 

DM: Now, they lived in the forest. They weren’t afraid of the forest? 

CJ: They were not afraid of anything. They were mostly—the men were armed with spears, but I never saw a 
machete in their hands either. They were armed with spears and they broke the plants apart to build their little 
huts and stuff; and they were totally forest people, they lived in the forest, they were like the forest animals. They 
just lived there. They were part of it. 

DM: Did they also use poison in the hunt? 

CJ: Yes. And they knew which it was; they knew what was poisonous and what was not. That’s evolutionary. 
That’s species selection right there. But they knew, and those several women and the two men were the only ones 
I ever saw, and I never saw any others. They went from camp to camp, but always within the big rainforest. They 


28 


Clyde Jones Memorial Volume 


never left it, and they never migrated, and they never saw the sea. They never saw anything other than—they 
were part of a forest—they were amazing people. 

DM: Even though the sea was how far from where they lived? 

CJ: Thirty miles, maybe. Yeah, maybe thirty miles... They never saw the sea or anything. And all of those 
women seemed to have a little age on them. I never saw young girls. I saw that one young boy. But I never saw 
any other young people. 

DM: I wonder why. 

CJ: I don’t know. 

DM: A high mortality rate, I imagine, but still— 

CJ: Child mortality was very high. But yeah, I never saw any in all the time I spent there. 

DM: Maybe they were afraid of you. Do you think they saw you, but kept their distance where you couldn’t 
see them? 

CJ: Sabater Pi talked about this, and he said they had them hidden somewhere. But I never saw any of them, 
other than that one young boy. 

DM: Which shows up in one of your photographs— 

CJ: Which is kind of tucked in behind me, hiding there— 

DM: He has kind of a western shirt on... 

CJ: I got that shirt for him; he didn’t have any clothes when I first saw him. He didn’t have any. I got those 
shorts and that shirt for him. I bought them and took them up there and helped him put them on, and I never 
saw him without them. He never took them off. And I never saw... they were very conscious about below their 
waist. They were totally opened above their waist but they were totally very conscious—the women were very 
conscious about the below the waist cover. 

DM: Were they carnivores, or did they eat the plants of the forest also? And did they seem to really know their 
botany? 

CJ: They knew the plants, especially those that were soft and those that they used for thatch and those that they 
used to build their huts. They ate very few fruits, they mostly were carnivores. 

DM: Okay, did you notice any medicinal use? 

CJ: They had some things. They would get some cuts and scratches. They had some plants they would rub 
on themselves, ostensibly for healing purposes. But, yeah, they ate—they had little nets that they spread in the 
forest and they caught dik-diks—the little antelopes—and they caught the giant flying squirrels that would come 
to the ground. Pangolins were a real treat for some reason. Pangolins were some kind of special treat for them. 


Marshall—Prologue 


29 


And then with their spears they took an occasional monkey. And they talked about killing an elephant for me, 
and I wouldn’t—they translated it back and forth, which took about half an hour to do anything. I wouldn’t let 
them and they acted put out about that. Most of these feelings were acts. They would act terribly put out about 
something. They were very fascinated with me writing my field notes. Every evening I would sit down and recap 
the day and write in my book. I had an audience every evening. 

DM: Did they seem to know what you were doing? Did they think this was some form of communication? 

CJ: Yeah. They were really attentive, but just ringed around me watching this every evening. It was an event. 
DM: Were they concerned about any predators in the forest? 

CJ: They didn’t seem to be. They didn’t seem to be concerned about the gorillas or the chimps which were 
almost right with them— 

DM: In size? 

CJ: Yeah. 

DM: Or larger, maybe? 

CJ: Larger. The adult gorilla and the adult chimp, they were larger, but I never saw them attack or try to kill one 
of those. They always took monkeys, and smaller ones. I never saw them mess with gorillas or chimps. They 
always—when a group of gorillas or chimps got near their little group of huts, they would yell and wave their 
hands and shoo them away. But I was very fascinated with that relationship. 

DM: Were there any predators in that area, like large cats of any kind that they should be concerned with, or 
would be concerned with? 

CJ: Yeah, there were West African lions, bush lions. And the main concerns were the elephants. They were 
concerned that the elephants would come through the camp and trash everything. We were out looking for gorillas 
and chimps once, and there were elephants nearby, and they indicated for me to climb into a small tree, so I did. 
But then I saw the elephants pushing trees over and I thought, “This is not a good idea.” And I got the heck out 
of there. That was their first response, climb a tree. No, no, that is not a good idea. They’re fascinating people. 
I think they were on the decline. Well, among the Fang, infant mortality initially was very high, and towards the 
end of my stay there, groups of missionaries came in and introduced baby formula. Infant mortality went way 
down. But everything started going to heck because, before, the women would nurse children until they were 
three and four years old. I mean I was sort of taken aback when a woman would be sitting in a chair or a stool 
and a youngster would walk up and stand there and start nursing, just everyday life. But infant mortality was 
increased and everything just sort of broke down at the end. The Guardia Civil thought they were very much 
in charge. But, you know, when they came to a village then everybody pretty much stood up, and they spoke to 
them and then they left and then they all kind of chatted, “Well, they’re gone now, no problem, we’ll go back to 
our old ways.” 

DM: It was a momentary disruption. 

CJ: Yeah. It was a strictly—a multicultural situation. Most of the country was occupied by the Fang, who were 
oriented toward the village and the slash-and-burn fields, with the hunter, who supplied bush meat. And there was 


30 


Clyde Jones Memorial Volume 


a group that lived along the coast called the Bubis. They were fishermen. They never went inland and there’s 
no intermixing between the two groups. And then the pygmoids. There was a Guardia Civil Spanish army, and 
then they had a local army that they were training. These were the young men that were supposed to be in train¬ 
ing to be in control.... 

They had a small navy. All they had were dugout canoes. 

DM: Did [the Bubis] use boats, did they use dugouts? 

CJ: Yeah. They went out and put out nets, and then they dried fish, and sold them to people. The Fang would 
come into Bata for the market. And then a few of the young men came in and had jobs, like, we had to hire a 
houseboy. When I stopped in Spain... they said I had to do several things. I had to hire a house person for the 
house, and I had to hire somebody to help maintain the yard. I didn’t need anybody, but I had to not disrupt the 
local economy. And I was advised that a good tracker—I might pay one U.S. dollar or the equivalent of one 
U.S. dollar a day, no more. And that seemed to be an amazing fee, to pay that. But I would hire somebody in 
one village, and in another village twenty miles away. When I got ready to hire the guy, he would say, “Well, 
you paid that guy the equivalent of a dollar, so many pesetas, and that’s what I charge.” And how they knew—I 
mean—they knew. I thought it was drums or something. Drums were active at night and I thought it had to be 
drums. That’s how they must have known. 

DM: Were there really drums active at night? 

CJ: Yes, there were really drums active at night. 

DM: You were probably the subject of some conversation in the province. 

CJ: They just knew everything; they just seemed to know everything. That happened to me, and everywhere I 
was, they seemed to know. How they knew, I have no idea. But they knew. And they would give me guidance: 
“Okay, now you paid that guy.” And also, I would rent a house in the Fang village. I’d rent a house, and the 
first thing I would do was negotiate the price, which was always equivalent to less than a dollar a day; it was like 
so many pesetas, like it was twelve to fifteen pesetas per dollar, and twelve pesetas was the order. And I would 
have to hire a young boy in the village to keep the goats off of me. The damn goats were everywhere. To keep 
the goats from getting on my bed, in the car, they were everywhere. And I would have to hire—they had a main 
house, a decent house, and then they had a house out back where the women lived and did the cooking—and that 
is where you went to eat, the women would heat food, or wash clothes, all for a price, of course. But it seemed 
to be twelve to fifteen pesetas, that seemed to be the going rate for everything. 

DM: Well, was there not much stratification of society? Were there not the rich and the poor among the Fang, 
for example? 

CJ: The goats were animals of wealth. 

DM: Oh yeah. 

CJ: Yeah, the guy who had fifty goats was better off than the guy that had twenty... And when it came time to 
butcher a goat, or eat a goat, usually the guy that only had twenty had to furnish the goat. But the damn goats 
were just, they were everywhere— 


Marshall—Prologue 


31 


DM: They did real well in that rainforest. 

CJ: They certainly did. And they milked them. They milked the female goats and had goat milk, which is good 
for you. 

DM: Was goat meat a frequent dish or was it more of a ceremonial dish? 

CJ: Not frequent. 

DM: Did they have sacrifices? 

CJ: They had— 

DM: Religious sacrifices? 

CJ: Yes, I think they did. And they had the chiefs—the head of the village had multiple wives, of course. The 
other thing I had to do was meet the chief, and meet all the wives. He would have them lined up. I met all of 
them you know: “ Umbulo, Umbulo, Umbulo, Umbulo.” And that was the group that would wash my clothes for 
me. For some reason, one of them would volunteer, one of them somewhere in the rank. They had some kind of 
rank in the way they were lined up, and one of them would volunteer to wash my clothes for me... 

The women always lived in the house behind the house. The head house is where the men lived and they had a 
house out back where the women lived. 

DM: That was standard throughout, then? 

CJ: Yes, everywhere... And the women had their mats and their beds in this cook house, and they had a constant 
fire going there, and they cooked there and they ate there. They did everything there and that’s where the women 
and kids lived. The men lived in the men’s house—interesting cultural thing... 

DM: You said there was not much interaction between the Bubi and the Fang. But what about the pygmoids? 
How did they all interrelate? Was there interrelation? 

CJ: I never saw any of the Fang interact with the pygmoids.... I never saw any of the Fang—there was no inter¬ 
breeding or any interaction with the Bubis other than the sale of fish. That’s all I ever saw. And the market was 
a wonderful experience. I was interested in the bush meat and it was everywhere: chimp arms and legs, gorilla 
arms and legs, and young gorillas and monkeys, and stuff like that. 

DM: When you say arms and legs, was it the arm with the hair and everything? 

CJ: Yeah. They just cut it off and there was an arm. When I was first there, before I got my Land Rover, I was 
riding the bus, and that was a trip. And that was an interesting thing. First of all, early on they wouldn’t let me 
on. Well, most of them had never seen anything like me. And this bus went on into the interior. And I’d ride the 
bus and it occurred to me what was happening—a bus would pull up to a bus stop, and there would be a structure 
there with an arm, and monkeys with the tail looped up around their necks. [They] made a slit around the tail 
and [they’d] bring it up and put it around the neck like a suitcase. And the monkeys would be on this structure, 
and the women would lean out of the bus and pull the hair on the bellies to see if it was fresh or not, because 
there was a difference in price. 


32 


Clyde Jones Memorial Volume 


DM: If it came right out, then it had been there too long. 

CJ: Yeah. But there was a definite difference in price between the old ones and the new ones. Well, then they 
would bring these stinking damn monkeys into the bus. And sometimes they’d just put the tail of the monkey 
over their head, or they’d put the monkey on their back and get off the bus. That’s also where I learned about the 
relationships of distance with people. I would be sitting on a log, waiting for the bus, and a hunter would come 
over the hill and the bus would stop everywhere, but he would come up and sit right next to me... That was an 
interesting kind of thing. It was as if we’d known each other all of our lives, you know. He would come up and 
put down and stick his cross bow in the ground and lay his machete down, and lay his blow gun across his lap 
and he’d sit right there... I always had an audience; everywhere I went, I had an audience right there... 

DM: They didn’t touch? 

CJ: No, they were close. They never touched. 

DM: Did they shake hands? 

CJ: After some work and, oh, when they got paid they would shake hands. And they had this other damnable 
habit—when you give them something and they have to give you something. And so I would—well, the money 
transaction was a very interesting thing. The grant money was sent to Tulane. Arthur Riopelle was there to 
handle communications and handle fiscal operations, which was very important. Money went from Tulane to 
the Chase Manhattan Bank, was transferred from the Chase Manhattan Bank to the Bank of Spain in Madrid, 
and it was transferred from the bank in Madrid to the branch bank in Bata. And they had the attitude in Bata 
that these people could never handle anything bigger than a hundred peseta note. So I would get my money 
in kind of a loaf of bread-type structure. And I [would] just put it in the seat of the Land Rover and drive out. 
And then I would pay these guys who had worked for me for four or five or seven days. Well, I’d pay them and 
they’d have to give me something. Early on, they’d give me a big stalk of bananas, and I would drive home and 
the kids would go “Oh boy, oh boy, bananas!” Well, several months later it was, “Oh God, bananas.” But that 
was a damnable custom they had. You couldn’t give anybody anything, but they’d have to give you something, 
whether you wanted it or not. 

DM: Was it always food? 

CJ: No, sometimes I would get a little wood carving, or a cane, or wooden spear, or a little figurine or something. 
That’s how I got most of the stuff that I got. 

DM: When you had your loaf of money, or anytime, really, did children beg? Did they come to you for little 
handouts? 

CJ: No, they didn’t beg. 

DM: That was not part of the culture. 

CJ: No, I never had a—in Bata I had a couple of beggars it seems but out in the villages I never felt begged upon, 
never. They used everything. They stockpiled everything in the woman’s house. And over in the corner there 
was a little fence-like thing. And it was stuffed with rags, cans, bottles, a piece of paper, everything. I mean, if 
I threw something out, it hardly hit the ground. It was gone. It was put in this. They would use it, eventually. 


Marshall—Prologue 


33 


DM: You pointed out in the photographs that there was no litter along the roads because of this. 

CJ: It was terribly clean. The whole country was terribly clean. And you noticed there were very few cars and 
fuel. The Land Rover was a diesel. And the guy that imported it and sold it to me explained that the diesel was 
the thing. That was the thing. 

DM: That’s what buses ran on also? 

CJ: The buses ran on diesel. This is 1967 and ’68. I was paying the equivalent of five dollars a gallon for diesel 
and I was very happy to get it. The first priority went to the buses, and what was left over was what I got. All 
the cars were diesel. What cars there were, they were diesels. 

DM: How far into the bush could you drive the Land Rover? It seems like your limitations would be pretty great. 

CJ: Well, it was a four wheel drive, which I hardly ever used because other than the main road, there was no side 
road that went anywhere. You walked. 

DM: You had to chop a trail if you walked? 

CJ: Yeah. To go someplace away from the main road you had to hire a man to lead you, and you had to hire 
a woman to carry your stuff. And I learned that this was usually a woman who was infertile—couldn’t bear 
children. And she was a—I have one of their carry baskets in that room. I don’t know if you’ve seen it or not. 
It’s a big tall one with—they’d load that thing up, and they’d put a band around it and put it on her forehead and 
put it on her back. 

DM: A tumpline. 

CJ: And sometimes it would weigh a hundred pounds. And we’d have to help her to her feet, and she would 
just go, and never stop. 

DM: She was infertile, so they found a niche for her. 

CJ: She had a place in the system, but she never left the trail and the trails were—sometimes you’d cross logs 
and stuff like that—sometimes they were primitive. And that village I showed you and I said “Those were the 
tracks from my Land Rover,” I think that was probably the first vehicle that village had ever seen. And they came 
out and they just touched the Land Rover. They just petted it, you know. But early on I was advised; I mean, it 
just had a canvas top, and you couldn’t lock it, and I had my camera and binoculars, a loaf of bread of money, 
everything in there. And they told me to find a woman who would put a little spirit on the Land Rover. And she 
gave me a little leaf-ball about the size of a golf ball, and it was on a little cord, a little vine cord, and I hung that 
on the rear view mirror. I never, ever lost a thing out of the Land Rover. And I would leave the Land Rover, and 
we’d be out in the woods for a week. 

DM: This was protection by the spirits. 

CJ: Yes. This was a voodoo-type thing. 


DM: Do you know what kind of leaf it was by any chance? 


34 


Clyde Jones Memorial Volume 


CJ: I have no idea what was in the—I was told don’t ever open it. You hang it there and leave it there and it 
will protect the Land Rover. And it did. I never lost—I would leave a loaf of bread of money. I would leave 
everything and be out in the woods for a week, and I never lost anything. It was total protection. 

DM: That’s amazing. 

CJ: Very strange. 

DM: Did children wear amulets around their necks and things like that? 

CJ: Sometimes, yes. 

DM: Similar kind of thing, leaves and—? 

CJ: Yeah. Very interesting people, good people, I learned high regard for them. 

DM: You mentioned that coming into this village... that they told you to drive your Land Rover across a couple 
of logs to get it over a creek or a river. 

CJ: Get it over a small ravine. And I wasn’t going to do it. And the logs had moss and stuff on them, and they 
took a machete and roughed up the top and said, “Okay, now drive across.” And I did. If it had fallen down, I 
never would have gotten it out. But yeah, they did, they went “buk, buk, buk, buk,” and roughed up the top so I 
could drive it and have traction. Clever! 

DM: There wouldn’t be much need for bridges out in the remote areas. Did people ford, or did they use logs 
for foot paths usually? 

CJ: Oh yes, every trail I was on had a log crossing a small ravine or something, or small creek or something, 
every one... I learned to just cross them. 

DM: Without thinking about it, just go on across? 

CJ: Yeah just go. And I always had my pack, and my camera, and my binoculars, and my stuff, you know. 

DM: You were talking about the market earlier, and you were talking about that level of interaction between the 
Bubi and the Fang—just the trade. Was it mostly barter, or was it money exchange? 

CJ: No, it was barter, they had very little money. 

DM: When the Fang made an exchange with the the bush meat hunters— 

CJ: Bush meat for fish. 

DM: That’s what it was, bush meat for fish. How did they [the Fang villagers] acquire the bush meat from the 
[Fang] hunters? What would they have exchanged? 

CJ: I never really figured that out. The hunter seemed to have some special role in the society of the village. It 
was always kind of aside, kind of different, because he went into the forest. That was one difference. He was a 


Marshall—Prologue 


35 


brave man, who went into the forest, and he always had his crossbow and his blow gun and his machete within 
reach, always within reach, and that’s how I would locate him. He’d be standing over there, kind of on the fringe. 
The whole village would come and listen to me, or be around to watch me, and he’d be over on the side, a little 
bit. But I never saw them give him any exchange, or any gifts or anything. That was just his role—was to go 
out and capture something; a dik-dik, or monkey, or something. 

DM: Did the villages have their own little exchanges? There was a market in the main towns, but was there any 
kind of little exchange going on in the small villages that you saw? 

CJ: A very special event would be butchering a goat. And they would hang the damn goat up by its hind legs, 
and cut its throat, and they’d scoop out a little dish in the sand, and the blood would flow into this place in the 
sand, and it would congeal, and then they would slice it, and eat it. 

And that was really tough for me. I think it was the texture of the sand that was really tough, which—I had to 
do it. That was the ceremony of butchering a goat—well, you see this in Mexico sometimes too, the congealed 
blood. They call it blood pudding, or whatever they call it. 

DM: Which makes you wonder about the universality of something like that, when you hear about it in the 
Americas and Africa. Did you get the indication that this sacrifice of a goat was in any way connected to a spirit 
or deity, or was it just an event in itself as far as you could tell? 

CJ: As far as I could tell it was just, yeah, they just selected a goat and butchered it. It was sort of a ceremonial 
thing, collecting of the blood, eating of the blood, and then they would divide up the meat, you know. They would 
take the damn machete and divvy up the goat in some sort of hierarchical structure in the village. 

DM: It was a community thing at least. 

CJ: It was. Everybody watched. 

DM: Did everybody regardless of age and gender eat? 

CJ: When they divvied up the meat it was a free for all—women, kids, men, everybody feasted and they also 
would trade bush meat for the little clothing they had. The Fang were the same way [as the pygmoids]; they were 
conscious about below the waist coverage, but very unconscious about the top. 

DM: Did you see any indication of prostitution? 

CJ: There were, no, but I saw a couple of cases—usually one of the wives would have sex with somebody else, 
and they would cut off her breasts. I saw several cases of that. I had a tracker once that was missing these two 
fingers. They were just gone. 

DM: The index and the middle finger. 

CJ: And I just thought it was a machete accident or something. And one time I asked him about it, and he got 
real embarrassed, and he walked away. And he came back and said “Okay,” he said, “I’m an ex-thief.” That is 
what he explained and that was the deal, and the drill in the village. If a woman committed adultery they’d cut off 
one or more of her breasts. How they all kept from bleeding to death, well they—they cauterized them. They’d 
take a stick out of the fire and the guy would just be there and not show any emotion or anything. I never saw 


36 


Clyde Jones Memorial Volume 


them cut off the breast of a woman, but it looked like it’d been cauterized. Again, they were totally unconscious 
about the top. Here’s a woman with one breast and just a big scar and advertising who she was. 

DM: Did you ever see any indication of a judicial procedure where they had a council within the village that 
sat for a judgment? 

CJ: The chief—the village chief, they went to him, and he made these decisions. 

DM: So they would just go to his home? 

CJ: Yeah, he had a place where he would sit and his wives would be there, and they would make the pitch, and 
he made the decision. 

DM: Was he chief for life or was this a—? 

CJ: I think it was once a chief, always a chief, all the time I was there. I got one of the little wands he had. He 
wanted to loan me one of his wives, and finally it hit on me—the explanation I could use was that it was against 
my religion. And when we were getting ready to leave, one of the guys was rather insistent on giving us a damn 
pig. And I wouldn’t eat, if I knew about it I wouldn’t eat their pork, because of infestations. But finally I had to 
explain that it was against my religion. Finally, I hit on that explanation, and they accepted that... 

I can tell you a very interesting experience. We used to go to the soccer games and... my former wife would not 
only be the only white woman, she’d be the only woman in the stands... Me and my wife and two kids—a little 
girl, and a little boy—had a place where they would put us. And we would go to the soccer games because there 
wasn’t anything else. And one time, one of my trackers came to my house and said, “You don’t want to go to 
the soccer game tomorrow.” And I thought, “Oh, he wants my tickets or something.” And he was very insistent: 
“No, you don’t want to go tomorrow. Or, if you go, don’t take the woman or the children.” So I took him. And 
so at intermission they brought out some bad guys, and killed them. And he knew that was going to happen, but 
he wouldn’t tell me what it was. He just said, “If you insist on going, don’t take the woman and children.” But 
that’s what they did. 

DM: How did they kill them? 

CJ: The Guardia Civil lined them up and just shot them. And I never knew what their crimes were. But, they just 
brought them out, and lined them up, and mowed them down, and dragged them off. It was just the thing to do. 

DM: As long as they had everybody together, there was a public execution. 

CJ: There was a lesson there, some kind of lesson. 

DM: These teams that would compete in soccer games, were they all within one village and divided into a team? 

CJ: Some teams came in from Cameroon and some teams came in from Gabon. Cameroon to the north, and 
Gabon to the south, they had some come in from across the borders. 

DM: So they had little provincial or national teams that moved about. 


Marshall—Prologue 


37 


CJ: Oh yeah, they were very strong, they had a very strong fan following. 

DM: Were they professional you think, or were they semiprofessional? 

CJ: I think they were semiprofessional; they were pretty good. 

DM: Another thing I wanted to ask you about the marketplace—we were talking about the Fang and the Bubi. 
Did the pygmoids ever come to the market? 

CJ: No. 

DM: Did they have any transactions at all outside of their group? 

CJ: Not that I ever saw. 

DM: Did they have any transactions within their group that you ever saw? 

CJ: Not that I ever saw; and they seemed to be—well, I described them as territorial; they seemed to just have 
a home range in which they worked, and that was theirs, and other groups were over there. I never saw any 
interactions with anybody. 

DM: They were completely self-sufficient, seems like. 

CJ: Totally. And where they got their clothes—I know they got them somewhere and that’s all they had, and 
they obviously never took them off. 

DM: They had cloth, didn’t they? Some of these [in the photographs] were made from bolts of cloth, it looks like. 

CJ: Well, the woman that adopted me just had one. She tied it around her waist, and that was all she had. And 
obviously they never took them off. 

DM: So they had a limited range and they lived completely off the land. Do you think they practiced infanticide, 
or did they even have to practice infanticide [with the high mortality rate]? 

CJ: That could have been one of the explanations of why I never saw young people other than this one. Maybe 
it was infanticide, maybe it was a limitation on what they could provide, because they knew within their area, 
they knew where all the animals were, they just knew. I mean, how they knew, I don’t know. It’s like they would 
have this conversation, and the two guys would have this conversation with the women. They’d have this group 
blab—a group conversation—and then a guy would explain, finally explain to me, that they were going to go get 
a monkey and they did. They knew where it was and they would go get it. Uncanny little animals, but really 
nice, after the initial standoffish bit with me, yeah, we became friends, and it was like they sort of took care of me. 

DM: Did they have any agriculture on any level? 

CJ: No. 


DM: Not even incipient? 


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CJ: None. The pygmoids had no agriculture at all. 

DM: Did they have any animal domestication? 

CJ: No. None. I never saw any. 

DM: Wow, the opportunity to see a culture like that! 

CJ: I would really like to know today, what’s happened to them. But I can’t go there; no white man can go to 
Equatorial Guinea. You chance your life if you go. First of all, you couldn’t get passage there; you’d have to 
walk in from Cameroon or Gabon, and you’d probably be killed. The word is out from the dictator. I know 
there was one Spanish guy that went in. When I was still teaching at Tech, Robert Owen was still there and there 
was a Spanish guy that worked on bats, and he came to spend some time with Owen. And he and I immediately 
became friends, because he had been back to Rio Muni, and that’s when he told me that there was no municipal 
power in Bata or any of the headquarter villages, there was no municipal power. The dictator had all this control, 
and it all went up to him. And being in the rainforest they had discovered oil, of course, and the oil companies 
were pouring—the figure is four million a year into it. And it all went to the dictator. The cultural salary and 
everything was still the same, according to him. They would say, “Yeah there was this guy here years ago, and 
he paid us twelve pesetas a day, and that’s what you pay us.” They knew and they remembered me, these various 
hunters remembered me. 

DM: So Robert Owen conveyed that information—that they still remembered your time there. 

CJ: And he introduced me to this guy and we became good friends, and he sat right here and explained modern 
day Rio Muni to me. I would like to go back, but on the other hand, I wouldn’t want to go back. My experience 
and my career are to go in and have your time, and you don’t go back. And that’s even becoming one of my 
thoughts about working in Big Bend and areas like that. For example, I wouldn’t go stay in Fajitas anymore. 
And that used to be a place where we always went and stayed, but I wouldn’t go there anymore. 

DM: You had your time. 

CJ: It’s different. 

DM: Right, a little bit heartbreaking sometimes? 

CJ: Yes. It’s very expensive, very different, and they closed the border, you know, the border patrol closed the 
border. 

DM: You can’t just wander across the border to Mexico. 

CJ: We used to just either walk across the border or drive over. I had a really wonderful experience, [when] 
we were walking across. I mean, the water hardly went above our ankles in the river, the Rio Grande, and we 
walked across. And we were walking up the sandy road and I found a marble just lying in the dirt. I found a 
marble. And there were a couple little kids playing marbles, and I got down and took a couple of shots and of 
course gave it to them. In a subsequent visit they came running up to me saying “Tengo canicas. Quiero jugar?” 
“I have marbles, do you want to play?” They remembered; they knew. 

DM: Did you ever see anything in Mexico—you did a lot of work in Mexico—did you ever see anything there 
that reminded you of Rio Muni? 


Marshall—Prologue 


39 


CJ: That behavior did. They knew, and in Mexico too it was the same thing, from one village to another, they 
knew we were coming, they knew what we were after. They knew we were after the rats and the bats, but they 
knew. And how in the hell they knew, I don’t know—yeah, they knew. That was the most interesting thing to 
me, was the communication that somehow preceded us. Didn’t you find that in the Tarahumara too? That they 
knew you were coming and they knew what you wanted to do? How the hell did they know? 

DM: It’s big news, I guess. 

CJ: Yeah, I don’t know how. 

DM: How it spreads, I’m not sure. 

CJ: Well, the one other thing about Mexico was they were always very sensitive. They would always say, “You 
can’t drink the water.” That was the first piece of advice we always—“You can’t drink the water; here you have 
to drink beer.” And their women and kids, little kids, they were drinking beer for breakfast, you know. But yeah, 
“You guys can’t drink the water; don’t drink the water.” Mike Bogan and I were working in Baja California Sur, 
and we stopped at a little truck stop. There were a couple of trucks parked there, so we stopped and the lady had 
a pot, and she was serving coffee to the truck drivers and we said, “Well, we’ll have a cup of coffee.” And she 
went to take the cup and dip it and the truck driver said “No,” and he explained to her, “ They can’t drink it like 
that.” So she took her apron and wiped out the cup, and gave it to us—cleaned it. 

DM: What were the water conditions in Rio Muni? You had a picture of drinking from a vine— 

CJ: Yeah, they would cut a piece of vine. They never carried water. They would always just cut a vine and drink 
it and discard the vine, or lay it by the trail, because it might be used later. The vine might be made into a blow 
gun even though it was crooked. It might be made into a blow gun and they would compensate for that, you know. 

DM: By aiming a little to one side or the other, (laughter) 

CJ: Yes, they would aim over here and they would shoot over there. They were uncanny in doing that. But 
yeah, the waters in the streams that were fast-moving cataract-type streams, that was—for them—that was safe 
water. They caught on very quickly that I wasn’t supposed to drink the water. I had to carry my own, which was 
bottled water from Bata. They had a remarkable little plant about the size of this room [about 144 square feet] 
in Bata, and it was a Coca-Cola plant and they had bottled water, which I had. 

DM: So you could drink that and you could drink from the vines. Do the vines affect you? 

CJ: No, but you know— 

DM: It was filtered enough? 

CJ: It was clean for some reason. 

DM: But they couldn’t just dip into a river. They had to get it also from a running water source, I mean a cascade? 

CJ: Yeah. But they knew what was safe water, and what wasn’t safe water. They knew. 

DM: Another interesting thing in those photographs was the huts that the pygmoids lived in. Can you describe them, 
and how they were constructed; what they used to cover them? It looked like maybe palm fronds or something. 


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Clyde Jones Memorial Volume 


CJ: All the roofs were thatched with palm fronds... In some cases, the walls were made of a series of small 
sticks and then packed with mud. 

DM: And were these the Fang or the pygmoids? 

CJ: The Fang. The pygmoids, they just had sticks. They had fronds and sticks and poles, small poles, and they 
would cut it up the center and off to the side, and they could make one of those huts while you watched. They 
could just make one. But they knew which plants to get, and they knew what poles to get. They knew everything. 

DM: Did you go inside these huts? Did they have any kind of baskets inside or storage of any kind? 

CJ: They’d have maybe a basket or two, or maybe, yeah, no pots... 

DM: Did they have any kind of matting for a bed? 

CJ: They would have a mat for a bed. They were really taken with my cot. I had a cot where you put the legs 
in and you would set it up about that high. They were really fascinated with that, and toward the end of my stay 
there, I gave my cot to one of my hunter friends Pancrasio Sima. I gave him my cot. Well, and the next place I 
went, they wanted to know why I didn’t give them my cot. 

DM: You disrupted the economy, (laughter) 

CJ: Yeah. There I went. 

DM: Hard not to do. So, okay— 

CJ: Long pants and long sleeve shirts... were a necessity because of the insects. And I would have a ring, just 
a raw ring around where the biting flies were on my wrist. When I was taking pictures or doing something, they 
would just have a ring around the end of my cuff. 

DM: You’ve given a good description about this before: It’s one hundred percent humidity, and I don’t know 
what the temperature is— 

CJ: Two hundred inches of rain a year—very high humidity. The temperatures were—when it got in the mid¬ 
seventies I was cold. And I think—because of the humidity—I was cold and I needed a blanket. 

DM: How warm was it when you went into the jungle? 

CJ: The high [was] 79, 80. 

DM: And you were buttoned up at the neck and the wrist, long sleeved— 

CJ: Long pants, long sleeves— 

DM: Because of the flies mostly— 

CJ: Insects, biting flies, and sometimes ants, soldier ants. I’d get into a pile of ants—well, then you’d just take 
your clothes off and get them off of you, [then] put your clothes back on, you know, no problem. 


Marshall—Prologue 


41 


DM: But you mentioned the ring of bites around your wrist, where the sweat was coming out. 

CJ: Yeah, the biting flies were going for moisture. And when it rained—well, my son wanted a rain gauge, so 
I had a rain gauge. Well, there came a shower, and the rain gauge overflowed. Well it would come up a shower 
and it would rain maybe seven inches, you know. It would just pour. And they [the locals] would just cut a leaf, 
and hold it over their head, and go on about their business. Y ou know, that was just the way it was. It rained—so 
it rained. No problem. 

DM: You said you got a touch of malaria. 

CJ: Yes, we all had malaria. My former wife and the kids had malaria. They remember being sick. They just— 
yesterday I was talking with my daughter and I mentioned what I was doing, and what you and I were doing, and 
she said she remembered having malaria. And then I had dengue fever. And I was out in the village and I got, 
I started to get “not myself.” I got sick. And my tracker insisted on riding back to Bata with me. But, well, he 
couldn’t drive. And we hit a road stop by the Guardia Civil. And they were mostly interested that I would let 
him in the Land Rover with me. That was the big issue, it was an issue—why was he in the Land Rover with 
me? Because, “No, no; they walk. They don’t ride, they walk.” And then I finally got it across that I was very 
sick and we got to Bata and I was really, really sick. And we went to see a Spanish doctor and he said “Well, I 
don’t know, come by tomorrow, maybe we can make a diagnosis.” I got up the next morning and I just had a 
red rash all over me. And he said “Oh we have a diagnosis. You have Dengue fever. Now, you probably won’t 
die of it.” Because I was a white man, yeah, probably won’t die of it. The Spanish doctors, I don’t know how 
they selected them, or how they chose to go to the colony, but if you ever needed a supply of gin you would find 
a Spanish doctor. He always had a supply. 

DM: For medicinal purposes? 

CJ: Of course, it scares off malaria. 

DM: Can you talk a little bit about your work there, tracking the lowland gorilla groups, for example? And 
especially relate the incident where you had a close call with a silverback. 

CJ: Well we would find a—my hunter-tracker and I—I had to pay him, because it took him from his job of col¬ 
lecting bush meat. I didn’t have to pay him for doing a service to me. I had to pay him for that distraction. So 
we would find a troupe of gorillas and we would follow them. And I would make observations and do whatever 
I did. And late in the study when the gorillas bedded down at night, I would bed down also. The tracker would 
make a little bed of vegetation for me, and I would stay awake and observe all night long. And that’s when I 
discovered that they did that [pats stomach rhythmically] all night long, lying on their back— 

DM: Patted their stomachs? 

CJ: Yes 

DM: And you could hear this? 

CJ: Yeah. And then finally, I figured it out, it was a form of communication. It was like “I’m okay. I’m over 
here and I’m okay. Are you okay?” That’s what it was, yeah. Jane Goodall agreed with me, that’s what it was... 

They had little fat bellies and they’re lying on their back [pats stomach]. A little drum thing like that would go 
on all night long; and I could hear that. “What the hell is that?” And that’s what it was. Well, then the next 


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Clyde Jones Memorial Volume 


morning the tracker would—well he would leave at night. He wouldn’t stay in the woods. He would leave, and 
the next morning his job was to come and find me; and he always did. He never ever left me alone [otherwise]. 
He never left me, or I wouldn’t be here. But he always came. And we were following a group of gorillas one 
time, and one of the showers had started, pouring down rain, and there was a big log that was down. And so we 
got under the log and we were sitting under the log, and I was writing notes. So we were under this log and we 
were whispering of course, which is what we always did. We whispered or communicated with hand signals. It 
stopped raining and I stood up and put my hands on this log. And there was like, from you to me, there was a 
silverback gorilla— 

DM: This is three feet away? 

CJ: And just like that, the other side of the log, like the distance from you to me. And he went, “Rawr! ” [shouted 
loudly]. And he spit seeds and stuff all over me, and he defecated all over my boots. And I’m not sure he was the 
only one [that defecated on the boots]. But that’s the closest I’ve ever been. And then he left; he didn’t attack; 
he left; he went. He screamed at me and then he left; he thundered off. And his females were following him. 
That was the closest I ever came and it was like a distance from you to me. 

DM: He was as startled as you? 

CJ: I guess he knew I was there... He screamed and spewed seeds —Aframomum seeds—all over me. 

DM: How long did it take you to recover from that? 

CJ: I sat back down and the tracker was lying on the ground going, “Pobre nosotros, pobre nosotros ”—“poor 
us, poor us, poor us.” 

DM: You said that sometimes—as I recall—sometimes they would get a little frustrated with your presence and 
break branches or trunks—small trunks. 

CJ: They would break branches, chest-beat, defecate, urinate, and go around and break stuff and throw branches, 
and throw defecate. And the gorillas would do that. And the real behaviorists were the chimps. And there’s 
a—Arthur Riopelle studied chimp distance. There’s a distance that you can get to a chimp and if you invade that 
distance, look out. And we did that once. We were following a group of chimps, and again it was raining, and 
we got too close and this big adult male chimp, about twice as big as me, came just thundering toward us. Just 
thundering, you know, breaking branches; just thundering toward us and grabbed a hold of a sapling about that 
big and just spun around it. 

DM: About four inches [in diameter]? 

CJ: Yeah, just spun around it just like a kid, spun around a pole. And we dove into the buttress of a tree. We 
were in there and my tracker was fooling around over there with something, cutting a pole, whacking something 
and it turned out to be a Gabon viper. And it just didn’t—it could have bitten both of us but it just didn’t. It’s 
one of those deals—it just didn’t. And I was watching this chimp. The chimp was just really creating a ruckus 
and the tracker was killing that viper. Yeah, it was one of those remarkable events; it just didn’t bite us. Because 
that’s the way they hunt. They just lay and they wait until something comes by. And that’s why they are colored 
the way they are. But that was a close call. 


DM: From two sides. 


Marshall—Prologue 


43 


CJ: I was more frightened of the chimp. I didn’t even see the damn snake but it was lying there in the leaves. 
DM: That’s as close as the chimp came, though—around the tree and then took off? 

CJ: Well, that was about from here to that wall. 

DM: That’s about ten feet. 

CJ: Yeah, and it looked—it spun around that tree, smashing everything, and hooting and hollering to beat hell, 
and throwing everything, throwing defecate, throwing everything. 

DM: So the beating of the chest was a warning. The beating of the tummy softly at night was, maybe, a sign 
of contentment. 

CJ: I’m convinced that was some kind of communication. And I’ve discussed that with Jane Goodall, and she 
agrees with me. She doesn’t know much about gorillas, but she knows a hell of a lot about chimps and she thinks 
it was a kind of communication. These little soft things like that. 

DM: What can you tell me about Jane Goodall’s personality? What is she like? What is her real contribution? 
Can you talk about her for a little bit? What was, maybe, your first impression? 

CJ: This was a young schoolgirl that Louis Leakey chose for some reason to go to Africa to study chimps. And 
she went to the Gombe Forest, and she took her mother with her the first trip so she wouldn’t die out there alone 
in the forest. But she learned how to work within the system and work with the native people in her area. And 
she spent twenty-five years studying chimps in the Gombe Stream area and the Gombe Stream Reserve—now it’s 
called. She’s written numerous papers on chimp behavior. She wrote a book summarizing her twenty-five years 
of studying chimps. And she’s a delightful person—quiet schoolgirl-type person. She was married once and had 
a child, reared the child in the Gombe Stream Reserve along with the chimps. And she and that husband divorced, 
and she has another husband that she lives with. He has a home in Mombasa. And she travels the world three 
hundred days a year giving talks in various places. She’s one of my favorite people. She’s a delightful person, very 
thoughtful, gave me a lot of good advice on how to deal with the local culture. Louis Leakey is now, of course, 
dead. But he had Jane Goodall, he had Dian Fossey, and there was another one that was studying highland goril¬ 
las. And she [Dian Fossey] was also a schoolgirl that he chose and supported her until she was murdered. And 
she was a delightful person also. She came to the National Geographic and came to the Smithsonian to visit with 
me several times, and we had lunch together several times. Louis Leakey had Jane Goodall, Dian Fossey, and I 
met him on my way to Rio Muni the first time. I stopped in Washington, D.C. and was at National Geographic. 
And Louis Leakey was there and we talked and had a really intense conversation for a couple of days. And then 
finally he said that “I like young people who don’t know anything.” And he liked me and he liked Jane Goodall, 
and he liked Dian Fossey. He liked young people who don’t know anything. 

DM: Was that distinction made at that point—with you working on lowland gorillas, Dian Fossey working on 
highland gorillas, and Jane Goodall working with chimps —Pan troglodytes , right? 

CJ: Yes, and I was the only one that was working with both gorillas and chimps. Dian Fossey was working 
with highland gorillas singularly, and Jane was working with chimps. I was the only one that had both of them 
in close sympatry and wrote a report on them, and wrote that report on the ecology of gorillas and chimps. Both 
Jane Goodall and Dian Fossey congratulated me on that piece of work. And you have Jane Goodall’s book on 
twenty-five years of chimp study and it has—inside the front cover is a small note from Jane. I reviewed that 


44 


Clyde Jones Memorial Volume 


book and she wrote a thank you note to me. And yeah, we were a far apart team—young people who don’t know 
anything. Jordi Sabater Pi came out of the research lab at the Barcelona Zoo, and for some reason Art Riopelle 
made contact with the director of the Barcelona Zoo, which was Senor Jonch, and he recommended that Sabater 
Pi would be our—we had to have a Spanish collaborator to go into a Spanish colony. Sabater Pi packed up his 
wife and two boys and moved to Bata, and he went out in the field with me on numerous occasions. He spoke 
some Fang and he was helpful. And later in time, he stayed in Bata, and he sort of did administrative stuff. He 
talked to the Guardia Civil. 

He was a Catalan. He was from Barcelona. He was a Catalan and he talked to people about me and what I was 
doing, and for them to leave me alone and he was very helpful in renewing—each of us had to have a passport. 
Two kids, and my wife, and me, each had to have a passport; and we could only have it for sixty days. And he 
was very helpful in renewing the passports in Rio Muni which was a problem every time. I got a guy, a Spanish 
guy yelling at me and complaining about me, and so I decided I’d just yell back. And so I said, “I’ll just take 
our passports and I’ll just leave.” And the guy just says, “Well,” and he just snatched one of the passports and he 
said, “We’ll just deport this one.” And it was my son’s passport and so I sat back down and we negotiated. And 
bribes were the order of the day. But late the next day—they had an immigration office manned by the Guardia 
Civil, and a couple of them standing around armed, and then they had an autonomous government immigration 
desk. And I would go there and say, “Well, I’ve dealt with the Guardia Civil” and they would say, “Well, we 
don’t care about that.” So bribes were the order of the day. I would have a loaf of bread and give them some 
money, both of them, the Spaniards and the Fang. But the civil governor of Rio Muni, for some reason he be¬ 
friended—he liked me for some reason and he and his wife, and me and my family, we would meet. They had a 
restaurant and a bar down on the beach, the beach bar, and we would meet there and have some drinks and some 
snacks. And that made us sort of standoffish too, because he was obviously befriending me. And the other thing 
that happened was, my God, the ambassador, the U.S. ambassador to Spain, came to the colony and came to see 
me, and, oh man, that did it. That set off the gossip that obviously I was a CIA officer, or some kind of spy, or 
something. They were really taken, that that guy would come to see me. And I was really taken by that too. I 
mean, here came his car with the flags on the fender and up my driveway and parked in front of my house, and I 
didn’t know he was coming. And he introduced himself as Angier Biddle Duke, the U.S. ambassador to Spain. 
And he had his wife with him and his daughter, and they were interested in what I was doing. They came in the 
house, after we recovered, and we told them what we were doing. 

DM: This was unannounced? 

CJ: Yes, just like that. I had some cages with some animals in them, and he wanted to see them. And they were 
really taken with the hairy frog and with the giant frog. And he was a very nice man, and she was, his wife was 
very nice. They were very common-type people. They came to see us. They came in our house. And, you know, 
we had, not lavish livings, just bare necessities. But those events really touched off both the Spaniards and the 
Fang. They were all both really taken by that and the Spaniards became very watchful of me after that. 

DM: And what about the Fang? 

CJ: Oh, they were just—they were kind of laughing about it. This high-powered person came to see me. 

DM: Now, Sabater Pi was there on behalf of the Barcelona Zoo? 

CJ: He was my coworker from Barcelona, Spain—from the Barcelona Zoo. And as it turned out later he—when 
he was stationed in Bata to do administrative things while I was out tromping around, he became an animal collec¬ 
tor, to collect animals and ship to the Barcelona Zoo, including the white gorilla. He got the famous white gorilla. 


Marshall—Prologue 


45 


DM: Named “Snowflake.” 

CJ: Yes. 

DM: The white gorilla. 

CJ: Copito de Nieve, Snowflake. 

DM: You visited years later in the Barcelona Zoo right? 

CJ: Yeah, it was a magnificent animal, huge, huge, male gorilla that sired numerous offspring. All of those were 
normal; all of those were black. 

DM: I’ve heard it said that this is the—not only a rare thing, an albino gorilla, but that Snowflake is the only 
known albino gorilla. Have you heard—? 

CJ: He’s the only one ever known, yeah, only one ever known. 

DM: I mention here for the recording: in the collection of photographs from S abater Pi, there are some of Snow¬ 
flake when he was a young gorilla, when he was first obtained. So that’s interesting as well. Well, what was his, 
Sabater Pi’s personality like? What kind of person was he? 

CJ: Well he was a Catalan, and one of our friends. One of the few other Spanish couples was a Basque, and he 
was the only other Spaniard there. And he was a fisherman, but he was the only other Spaniard, so naturally he 
and Sabater got together frequently. And there were always these political discussions about the Basques sepa¬ 
rating from Spain and the Catalans separating from Spain; and there was always this, every time, this political 
discussion. But Sabater Pi, he was a nice man, very set in his ways, and very stubborn. The typical Catalan, I 
think. He and his wife, and two sons, were friends of ours. They were good to us. He coauthored numerous 
papers with me including the gorilla-chimp one that you have [“Comparative Ecology of Gorilla gorilla and Pan 
troglodytes in Rio Muni”] but he was a little stubborn. 

DM: He was not an academically trained scientist, was he? 

CJ: No, he was just a person that rose up, like some in the U.S. that I got to know when I was at the Smithsonian 
who just, you know, started out as a squeegee person and later rose up to be a curator. He floated up due to his 
enthusiasm. He worked hard—he was a hard worker. 

DM: It wasn’t family connections or anything like that? 

CJ: No. 

DM: Did he coauthor with you the article on chimpanzee use of tools [“Sticks Used by Chimpanzees in Rio Muni”]? 
CJ: Yes, which you have, I believe. 

DM: Can you tell me how that research came about, and that discovery? 


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Clyde Jones Memorial Volume 


CJ: Well, we just discovered chimps hovered around a termite mound, and lots of hooting and hollering, lots of, 
you know, jumping around. And then we saw these sticks lying there. We just looked at the termite mound and 
there were these sticks, and holy cow! 

DM: Were they lying there or were they stuck into the mound? 

CJ: They were both stuck in and others just thrown down, and so I just collected a bunch of these sticks and we 
got to observing this. This is obviously a learned behavior trait, using these sticks to stick in the termite mound and 
pull out— there would be a termite attached to it, a Macrotermes muelleri, and they’re about that long, big ones— 

DM: About a quarter-inch or half-inch long? 

CJ: About a half-inch long, and about half of that was jaws. And the hooting and hollering was—they would 
bite the chimps on the lips. But they were eating them, because they were a good source of protein, but, yeah— 

DM: And, by the way, there are photographs of these termite mounds and sticks, also in your collection. 

CJ: I had a couple, you saw a couple that I had. 

DM: And there’s a sketch, also on the front, I believe, of the article, and I don’t know if that was a Sabater Pi 
sketch— 

CJ: No, that was a sketch made by Wilma Martin of the Delta Primate Center... Leonard Carmichael paid for 
me to be at the Primate Center for a year to write, to write the results of my work, and look for a job. 

DM: At about the same time, Jane Goodall was doing similar work. I don’t know if she was specifying the use 
of sticks in termite mounds— 

CJ: She got on to it. 

DM: Okay, was this independent of each other? 

CJ: No, we communicated that, and she got on to it. She found it and it was an interesting find, I thought. 

DM: Did your [discovery] precede hers, do you think? Or... did she hear about your research? 

CJ: Yes, we talked about it, and I recall talking to her about it. But she got on to it too about the same time. And 
I collected—those sticks are at Tulane. 

DM: That was my next question. And they are, hopefully, tagged and described. 

CJ: They are labeled as if they were specimens in the mammal collection at Tulane. The interesting thing is that 
[my] longtime friend and curator at Tulane is dead and they don’t have a mammalogist. A1 Gardner, who is at 
the National Museum at the Smithsonian, he was hired to replace me at Tulane and then I hired him, I hired him 
to come to D.C. And Tulane has never replaced him and they’ve never had a mammalogist since then. But my 
friend [Royal] Suttkus [a professor at Tulane]—who was a collector of everything—he took care of the mammals. 


Marshall—Prologue 


47 


DM: I know one of his big things was fish. 

CJ: Yeah, oh, yes, like several million fish. And that is the biggest collection of freshwater fishes in the world. 
Well anyway, so A1 Gardner goes down to Tulane—his daughter is a student at Tulane—A1 Gardner goes to New 
Orleans and he packs up the mammals that I got from Rio Muni and took them to the National Museum. He just 
packed them up, and took them to D.C. 

DM: Yet, the sticks are still there—? 

CJ: Yeah. 

DM: At Tulane? 

CJ: I was impressed, he just packed them up. 

DM: So to see this work, a lot of the work you did, you go to the National Museum [of Natural History], 

CJ: So all the flying squirrels, the pangolins, that stuff, that’s all now at the National Museum, where it should 
be, because they have an African initiative. 


[At present, the Clyde Jones specimen collection from Rio Muni is housed at the National Museum of Natural 
History, Washington, D.C.; the bulk of his Chihuahuan Desert specimen collection is at the Natural Science 
Research Laboratory, Museum of Texas Tech University; and his interviews, manuscripts, and photographs, 
including those appearing in this article, are at the Southwest Collection/Special Collections Library, Texas Tech 
University, Lubbock, Texas.] 


Works Cited 


Jones, Clyde. Interview by David Marshall and William Ty- 
deman, November 6,2003, in Lubbock, Texas. Compact 
disc. Southwest Collection/Special Collections Library, 
Texas Tech University, Lubbock. 


Jones, Clyde. Interview by David Marshall, February 27,2012, 
in Lubbock, Texas. Compact disc. Southwest Collec¬ 
tion/Special Collections Library, Texas Tech University, 
Lubbock. 


A New Species of Myotis (Chiroptera: Vespertilionidae) from Suriname 

Ricardo Moratelli, Don E. Wilson, Alfred L. Gardner, Robert D. Fisher, and Eliecer E. Gutierrez 


Abstract 

We describe a new species of bat in the genus Myotis (Vespertilionidae: Myotinae) 
from the district of Sipaliwini, Suriname. The new species ( Myotis clydejonesi sp. 
nov.), known from a single specimen, is sister to a clade of M. nigricans (Schinz) from 
southern South America, but differs from all Neotropical species of Myotis in qualita¬ 
tive and quantitative morphological characters and in its cytochrome-/? gene sequence. 
Our findings also indicate that M. nigricans remains composite and provide support for 
restricting M. nigricans (sensu stricto) to southern South America. 

Key words: Guiana Shield, Myotinae, Neotropics, South America 


Introduction 


The Guiana Shield comprises part of eastern Co¬ 
lombia, southern Venezuela, northern Brazil, Guyana, 
Suriname, and French Guiana (Gibbs and Barron 1993; 
Huber 1994). Some authors (e.g., Hollowed et al. 2001; 
Lim et al. 2005) exclude eastern Colombia and northern 
Brazil from their definition of the Guiana Shield. For 
this more restricted area, Lim et al. (2005) reported 
148 species of bats, including five species of Myotis 
Kaup, 1829— Myotis albescens (Geoffroy, 1806), M. 
nigricans (Schinz, 1821), M. oxyotus (Peters, 1866), 
M. keaysi Allen, 1914, and M. riparius Handley, 1960. 
Three of these ( albescens , nigricans, and riparius ) oc¬ 
cur in Suriname (Husson 1962; Lim et al. 2005). 

The three species that occur in Suriname are wide¬ 
spread in the Neotropics, their distributions extending 
from Central America southward into southern South 


America (Wilson 2008). Among them, M. albescens 
has been retrieved as a monophyletic, morphologically 
cohesive group (Moratelli and Oliveira 2011; Larsen et 
al. 2012). On the other hand, M. nigricans, as currently 
recognized, appears to be a composite of several spe¬ 
cies (Moratelli et al. 2011, 2013; Larsen et al. 2012). 

In the course of a critical review of collections 
of Neotropical Myotis , we found one specimen from 
Sipaliwini, Suriname, that has a peculiar cranial mor¬ 
phology. Based on pelage color and texture, and cranial 
features, this specimen is unquestionably allied with 
species in the albescens group (sensu Moratelli et al. 
2013), but qualitative and quantitative morphological 
features, along with its cytochrome-/? ( Cytb ) profile, 
distinguish the Suriname specimen from these and all 
other Neotropical Myotis. 


Methods 


Specimens examined. —The source of the mate¬ 
rial for the description of this new species is one adult 
lactating female deposited in the Museum of Texas 
Tech University (TTU 109227). It was collected by 
H. H. Genoways on 23 January 2008 at Raleigh Falls 
(04°43' N, 56° 12' W), district of Sipaliwini, Suriname. 


This research is part of a critical review of collec¬ 
tions of Neotropical Myotis for which more than 3,800 
specimens have been examined, including all species 
currently recognized (see Moratelli and Wilson 2014). 
Recognizing TTU 109227 as unusual, we compared it 
directly with 368 vouchers (Appendix) representing 


49 


50 


Clyde Jones Memorial Volume 


all species currently recognized from northern South 
America (see Moratelli et al. 2013), giving special at¬ 
tention to those species from the Guiana Shield (see 
Moratelli et al. 2015). These vouchers are preserved 
in the American Museum of Natural History (AMNH, 
New York, USA); Carnegie Museum of Natural His¬ 
tory (CM, Pittsburgh, USA); Centre for the Study 
of Biological Diversity, University of Guyana (M, 
Georgetown, Guyana); Museum d’histoire naturelle 
(MHNG, Geneva, Switzerland); Museum of Texas 
Tech University (TTU, Lubbock, USA); Museu de 
Zoologia da Universidade de Sao Paulo (MZUSP, Sao 
Paulo, Brazil); Natural History Museum, University 
of Kansas (KU, Lawrence, USA); National Museum 
of Natural History, Smithsonian Institution (USNM, 
Washington, DC, USA); and Royal Ontario Museum 
(ROM, Toronto, Canada). These specimens were 
identified according to criteria described by Moratelli 
et al. (2013,2015). 

Morphology and morphometries. —Descriptive 
terminology for craniodental morphology follows 
Moratelli et al. (2013). Measurements were taken of 
adults only, and are reported in millimeters (mm), and 
the body mass is in grams (g). We recorded the total 
length (TL), tail, hind foot, ear, and body mass from 
skin labels, reported to the nearest millimeter or gram. 
Other measurements were taken using digital calipers 
accurate to 0.02 mm. Craniodental measurements were 
taken with the aid of binocular microscopes under low 
magnification (usually 6x). These dimensions were re¬ 
corded and analyzed to the nearest 0.01 mm, but values 
were rounded off to 0.1 mm throughout the text because 
this is the smallest unit that allows accurate repeatabil¬ 
ity with calipers (Voss et al. 2013). Measurements, as 
defined in Moratelli et al. (2013:3), include forearm 
length (FA), third metacarpal length (3MC), length of 
dorsal hair (LDH), length of ventral hair (LVH), great¬ 
est length of skull (GLS), condylocanine length (CCL), 
condylobasal length (CBL), condyloincisive length 
(CIL), basal length (BAL), zygomatic breadth (ZB), 
mastoid breadth (MAB), braincase breadth (BCB), 
interorbital breadth (IOB), postorbital breadth (POB), 
breadth across canines (BAC), breadth across molars 
(BAM), maxillary toothrow length (MTL), length of 
the upper molars (Ml-3), mandibular length (MAL), 
and mandibular toothrow length (MAN). Descriptive 
statistics (mean and range) were calculated for all 


dimensions with sample size {ri)> 3. A discriminant 
function analysis (DFA) was applied to a subset of the 
craniodental dimensions (MAB, CIL, MAL, BAL, 
GLS, POB, Ml-3, BAC) to compare TTU 109227 with 
representatives of the most similar species. Statistics 
was performed in SPSS (IBM Corp. 2012). The list of 
specimens used in the DFA is in the Appendix. 

Phylogenetic analyses. —Phylogenetic analyses 
of Cytb sequences were conducted for Neotropical 
species of Myotis, which Ruedi et al. (2013) found to 
represent a monophyletic group. A total of 118 Cytb 
sequences for species in this clade, and four and seven 
sequences for Myotis brandtii and M. gracilis , respec¬ 
tively, were retrieved from GenBank (Table 1). We 
used Myotis brandtii and M. gracilis as an outgroup 
because Ruedi et al. (2013) found they were sister to 
the Neotropical clade. Sequences were aligned using 
default options of MAFFT v.7.017 (Katoh and Standley 
2013) as implemented in Geneious v.7.1.5 (Biomatters, 
http://www.geneious.com/). Subsequently, the Bayes¬ 
ian Information Criterion (BIC), as implemented in Par- 
titionFinder ver. 1.0.1 (Lanfear et al. 2012), was used 
to determine both the most suitable partition scheme 
and the best-fit models of nucleotide substitution. This 
analysis only considered models that can be applied in 
MrBayes (see below). 

Maximum likelihood (ML) and Bayesian infer¬ 
ence (BI) were used as optimality criteria. The ML 
analysis consisted of 20 independent searches in the 
Genetic Algorithm for Rapid Likelihood Inference 
(GARLI 2.0; Zwickl 2006) applying the best fit-model 
and the best partitioning scheme (see Results) and de¬ 
fault settings. The Bayesian analysis was conducted 
in MrBayes v. 3.2 (Ronquist et al. 2012). The search 
started with a random tree. The Markov chains were 
run for 100 million generations, and trees were sampled 
every 1,000 generations. Default values were kept for 
the “relburnin” and “burninfrac” options in MrBayes; 
therefore, the first 25,000,000 generations (25,000 
trees) were discarded as burn-in, and posterior prob¬ 
ability estimates of all model parameters were based 
on the remaining (75,000) trees. Convergence and 
stationarity were assessed in the Bayesian analyses by 
plotting likelihood values in Tracer 1.5 (Rambaut and 
Drummond 2007). 


Moratelli et al.—New Myotis from Suriname 


51 


Table 1. Terminals (focal species and putative species of the genus Myotis; see Methods) and corresponding GenBank 
accession numbers. Note that the information presented herein for terminal taxonomic identifications results from 
re-identification of voucher specimens (see Methods), and do not necessarily match those identifications assigned by 
researchers that generated the corresponding sequence(s) available at GenBank. 


Terminal 

GenBank accession number 

M. albescens 

AF376839, JX130444, JX130445, JX130463-JX130465, JX130472, 
JX130500-JX130504, JX130522 

M. atacamensis 

AM261882 

M. austroriparius 

AM261885 

M. brandtii 

AF376844, AM261886, AY665139, AY665168 

M. cf. lavali 

AF376864 

M. cf. nigricans (Suriname) 

JN020570-JN020572 

M. cf. nigricans (Tobago) 

JN020573, JN020574 

M. cf. nigricans (western Ecuador) 

JX130523, JX130541, JX130546-JX130550 

M. cf. nigricans (eastern Pem) 

JX130452, JX130537, JX130538 

M. cf. pilosatibialis 

AF376852, JX130449, JX130489, JX130514, JX130519, JX130525 

M. chiloensis 

AM261888 

M. clydejonesi 

JX130520 

M. dinellii 

JX130475 

M. dominicensis 

AF376848, JN020554-JN020556 

M. gracilis 

AB106609, AB243025-AB243030 

M. handleyi 

JN020569, JX130529-JX130533, JX130535, JX130543, JX130544 

M. levis 

AF376853 

M. martiniquensis 

AM262332, JN020557-JN020561 

Myotis sp. 

JX130493 

M. nesopolus 

JN020575-JN020577 

M. nigricans 

JX130450, JX130455, JX130496, JX130498, JX130499, JX130528, JX130539, 
JX130540 

M. nyctor 

JN020562-JN020567 

M. oxyotus 

AF376865 

M. pilosatibialis 

JX130526 

M. riparius 

AF376866, AF376867, AM261891, AM262336, JX130436, JX130469, 
JX130473, JX130474, JX130479-JX130481, JX130485, JX130486, JX130488, 
JX130491, JX130492, JX130506, JX130513, JX130515, JX130516, JX130572 

M. velifer 

AF376870, AY460343, EF222340, EU680298, EU680299, JX130438, 
JX130462, JX130468, JX130477, JX130478, JX130589, JX130592 

M. vivesi 

AJ504406, AJ504407 

M. yumanensis 

AF376875 


52 


Clyde Jones Memorial Volume 


Nonparametric bootstrapping (Felsenstein 1985) 
for the ML analysis, and posterior probabilities for the 
BI analysis, were used to assess nodal support (Ron- 
quist et al. 2012). The ML bootstrap analysis was per¬ 
formed in GARLI 2.0 using 100 pseudoreplicated data 
matrices, with 10 searches performed on each. Bayes¬ 
ian posterior probabilities were calculated simultane¬ 
ously with the search for the best Bayesian topology, 
conducted as described earlier. Throughout the text, 
we refer to different degrees of nodal support for the 
ML bootstrap analysis using the following categories: 
strong support, for bootstrap values >75%; moderate 
support, for bootstrap values > 50% and < 75%; negli¬ 
gible support, for values < 50%. For the BI analysis, we 
refer to degrees of nodal support with two categories, 


significant or strong in cases in which a node’s posterior 
probability was > 0.95, and insignificant or negligible 
for posterior probability values <0.95. 

High values of sequence divergences are neither 
necessary nor sufficient for recognition of linages at 
the species level (Ferguson 2002; Davalos and Russell 
2014). However, genetic distances provide a heuristi- 
cally useful basis for comparisons of genetic variation 
within and among lineages (Gutierrez et al. 2010). 
Therefore, we report average uncorrected (p) distance 
and average Kimura 2-parameter-corrected (K2P) dis¬ 
tance within and among haplogroups of interest for our 
taxonomic objective. Genetic distances were calculated 
using MEGA version 5.2.1 (Tamura et al. 2011). 


Results 


Molecular analyses. —The Cytb matrix contained 
ca. 8% of missing data. PartitionFinder found that the 
most suitable partitioning scheme was not to use sub¬ 
sets, and that the best-fit model of nucleotide substitu¬ 
tion was the Hasegawa, Kishino, and Yano model, with 
gamma-distributed rate heterogeneity and a proportion 
of invariant sites (HKY + T+ I). 

Our results show that M. nigricans , as currently 
understood, is polyphyletic, with representatives in 
five distinct, strongly supported haplogroups (Fig. 1). 
One comprises samples from Bolivia and Paraguay, 
and whose sister terminal was found to be specimen 
TTU 109227. Another comprises samples from eastern 
Peru, and was recovered sister to M. nesopolus Miller, 
1900, albeit with negligible nodal support. The third 
group comprises samples from western Ecuador, but 
its relationship to other closely related haplogroups 
remains equivocal. The fourth includes two samples 
from Tobago and was recovered, with strong support, 
as sister to the clade including the M. cf. nigricans 
haplogroup from western Ecuador; M. cf. handleyi, 
Myotis sp., M. cf. nigricans from eastern Peru; and 
M. nesopolus. The last M. cf. nigricans haplogroup 
includes samples from Suriname (except TTU 109227), 
and was recovered, with strong support, as sister to 
M. nyctor LaVal and Schwartz, 1974 from the Lesser 
Antilles. Our primary motivation for this research 
was to determine the identity and taxonomic status 


of TTU 109227; hence, our results and discussion are 
focused on the phylogenetic positioning and morpho¬ 
logical distinctiveness of this specimen. From here 
on we refer to Bolivian and Paraguayan samples as 
M. nigricans due to geographical proximity to its type 
locality (southeastern Brazil [LaVal 1973; Moratelli et 
al. 2011]). We refer to the other population samples in 
the remaining haplogroups tentatively assigned to M. 
nigricans as M. cf. nigricans. 

We found the specimen of interest—TTU 
109227—to be sister to M. nigricans (Fig. 1). Sequence 
divergences between TTU 109227 and M. nigricans 
are 5.4% and 5.7% for p- and K2P-distances, respec¬ 
tively. Note that to accomplish the calculation of these 
between-groups distances, the DNA sequence of speci¬ 
men TTU 109227 was duplicated. The within-group 
sequence divergence for the M. nigricans haplogroup 
is 1.5% for both metrics. Within-group sequence 
divergence could not be calculated for the clade con¬ 
taining TTU 109227 because only a single sequence 
is available for it. 

Morphological analysis. —Considering the 
assemblages of Myotis known from northern South 
America (Moratelli et al. 2013) and the Guiana Shield 
(Moratelli et al. 2015), the Surinamese voucher TTU 
109227 can be distinguished morphologically from 
species in the ruber group (i.e., keaysi, pilosatibialis, 


Moratelli et al.—New Myotis from Suriname 


53 


0.00012 substitutions per site 

Figure 1. Phylogenetic tree resulting from the Bayesian inference analysis of the 
cytochrome-/? sequence data. Nodal support from both the Bayesian inference and the 
maximum-likelihood analyses are shown right and left of slashes (“/”), respectively. See 
Methods for more information. 


54 


Clyde Jones Memorial Volume 


riparius, and simus) by its silky pelage, low sagittal and 
lambdoidal crests, and rounded, relatively uninflated 
occipital region. TTU 109227 is morphologically 
close to species in the albescens group (i.e., albescens, 
caucensis, handleyi, nigricans, oxyotus, and nesopolus 
larensis ) in the traits described above. However, it can 
be distinguished from all species in this group by the 
depressed braincase (Fig. 2), and the unique combina¬ 
tion of dorsal fur blackish and ventral fur with blackish 
bases and yellowish-red tips. FromM nigricans —the 
most closely related species—TTU 109227 can be 
distinguished by its anteriorly shallower rostrum, more 
robust occipital condyles, supraoccipital not as inflated 
and not projecting as far behind occipital condyles (Fig. 
2), and more laterally expanded mastoid region. We 


provide additional information on its description and 
distinction from other species under the subheading 
“Morphological description and comparisons.” 

Morphometric analyses. —In a discriminant 
analysis, the cranial morphology of TTU 109227 was 
compared with the morphology of M. nigricans from 
Bolivia and Paraguay and of M. cf. nigricans from 
Suriname (Fig. 3, Table 2). In this analysis, the first 
two discriminant functions (DF1 and DF2) summarized 
100% of the among-group variation, with DF1 com¬ 
prising 82% and DF2 18%. Along the first axis (DF 1), 
TTU 109227 andM nigricans had low negative values, 
and M. cf. nigricans had high positive values. Along 
the second axis (DF2), TTU 109227 had high negative 


Figure 2. Lateral (A), ventral (B), and dorsal (C) views of the skull of the holotype of Myotis clydejonesi 
(TTU 109227), and lateral (D), ventral (E), and dorsal (F) views of the neotype of M. nigricans (LACM 
36877). Scale bar = 5 mm. See Table 1 for measurements. Arrows indicate the comparatively depressed 
braincase (1), anteriorly shallower rostrum (2), more robust occipital condyles (3), and more laterally 
expanded mastoid region (4). 


Moratelli et al.—New Myotis from Suriname 


55 


Discriminant Function 1 (82%) 

Figure 3. Plots of multivariate individual scores in the first two 
discriminant functions (DF1, DF2). Samples: M. nigricans 
from Paraguay (n = 10 [circles; group 1]), M. cf. nigricans 
from Suriname (n- 3 [diamonds; group 2]), and the holotype 
of Myotis clydejonesi (triangle; group 3). 


Table 2. Coefficients of discriminant functions (DF1, DF2) for samples 
of Myotis clydejonesi, M. cf. nigricans, and M. nigricans. See Methods 
for variable abbreviations. 


DF1 

DF2 

Characters 

82.4% 

17.6% 

GLS 

-2.311 

3.359 

CIL 

-25.173 

1.013 

BAL 

10.071 

-4.457 

MAB 

1.353 

1.327 

POB 

3.272 

0.711 

BAC 

-4.972 

3.782 

Ml-3 

4.049 

0.423 

MAL 

18.634 

-4.209 


56 


Clyde Jones Memorial Volume 


values, whereas M. nigricans and M. cf. nigricans had 
low negative to high positive values. Thus, this analysis 
confirmed the distinctive cranial morphology peculiar 
to TTU 109227. 

Comparing linear measurements of TTU 109227 
with M. nigricans (n = 41-54) and M. cf. nigricans (n 
= 2-3), TTU 109227 has larger dimensions, without 
overlap, for most characters (all but Ml-3) related to 
the length of skull (GLS, CCL, CBL, CIL, BAL) and 
rostrum (MTL, Ml-3, MAL, MAN). On the other 
hand, all width measurements (MAB, BCB, IOB, POB) 
and rostrum (BAC, BAM) overlap with those from M. 
nigricans and M. cf. nigricans. These results indicate 
that TTU 109227 has a skull comparatively longer, 
but not wider, than the skulls of M. cf. nigricans and 
M. nigricans. 

Combined results from morphological, morpho¬ 
metric, and molecular analyses show that TTU 109227 
represents a unique lineage that differs from all species 
of Neotropical Myotis in qualitative and quantitative 
morphological characters and in its Cytb gene sequence. 
Based on these findings, we recognize TTU 109227 as 
a representative of an undescribed species, which we 
here name as: 

Myotis clydejonesi sp. nov. 

Clyde Jones’s Myotis, Myotis de Clyde Jones 
Figs. 2, 4, 5; Table 3 

Holotype and type locality. —The holotype (TTU 
109227) comprises the skin and skull of an adult lactat- 
ing female (Figs. 4, 5), including tissue (TK 151465), 
collected by H. H. Genoways (field number 6630) on 
23 January 2008 at Raleigh Falls (04°43' N, 56° 12' W; 
obtained from the skin label), Sipaliwini, Suriname. 
External and craniodental dimensions are in Table 3. 
The species is known from only the type locality (Fig. 
6). This collecting site is located on an island in the 
Coppername River in the Central Suriname Nature Re¬ 
serve. Relatively dense, near-mature tropical lowland 
forest with only a limited understory occupied most 
of the area. The mist net in which the holotype was 
captured was placed under the largest tropical trees 
near the banks of the Coppername River (Genoways 
and McLaren 2003). 


Diagnosis.—Myotis clydejonesi can be distin¬ 
guished from all other Neotropical species of Myotis 
by the flattened braincase, elongated rostrum, silky 
fur, and combination of dorsal and ventral pelage 
colors. The fur is silky; dorsal pelage is blackish, 
without contrast between bases and tips, and ventral 
fur is blackish basally (2/3 of the total hair length) and 
tipped yellowish-red on terminal third. The braincase 
is flatter and the pelage is silkier than in any other South 
American species known to us. This combination of the 
ventral and dorsal pelage colors appears to be unique 
among Neotropical Myotis. The following set of traits 
also is useful to distinguish M. clydejonesi from other 
Myotis that occur on the Guiana Shield: long, silky 
pelage; absence of a fringe along the trailing edge of 
uropatagium; low sagittal and lambdoidal crests; and 
rounded occipital region. 

Morphological description and comparisons .— 
Among South American Myotis, M. clydejonesi is a 
medium-sized species (FA 34.9 mm, other measure¬ 
ments in Table 3). The pelage is silky. Dorsal fur 
is blackish without contrast between bases and tips. 
Ventral fur is blackish basally (2/3 of the total fur 
length) and yellowish-red on the tips (1/3), with strong 
contrast in color between bases and tips. Membranes 
are medium-brown. The plagiopatagium is attached 
to the foot at the level of the toes by a broad band of 
membrane (see Lopez-Gonzalez et al. 2001:141, fig. 
la). The dorsal surfaces of elbow and tibia are naked or 
nearly naked. The uropatagium lacks the fringe of hairs 
along the trailing edge. Like most species of Myotis, 
its dental formula is 2/3, 1/1, 3/3, 3/3 = 38. The P3 is 
aligned in the toothrow (not displaced lingually), and 
visible in lateral view. Frontals are slightly inclined, 
with a smooth transition from the rostrum to the brain¬ 
case. The sagittal and lambdoidal crests are low. The 
occipital region is rounded, and does not project much 
behind level of occipital condyles. In contrast to other 
Neotropical Myotis, the skull is flattened, but not nearly 
so flattened as in M. planiceps Baker, 1955 (see photo of 
aM. planiceps skull in Haynie et al. 2016:703, fig. 2a). 

In addition to the diagnostic traits, M. clydejonesi 
can be distinguished from those species that co-occur 
on the Guiana Shield ( albescens, keaysi, riparius, 
oxyotus, nigricans) as follows: from M. albescens by 


Moratelli et al.—New Myotis from Suriname 


57 


Figure 4. Dorsal (A) and ventral (B) views of the skin of the holotype of Myotis 
clydejonesi (TTU 109227). See Table 3 for measurements. 


Figure 5. Dorsal (A), ventral (B), and 
lateral (D) views of the skull, and dorsal 
(C) and lateral (E) views of the mandible 
of the holotype of Myotis clydejonesi 
(TTU 109227). Scale bar = 5 mm. See 
Table 3 for measurements. 


58 


Clyde Jones Memorial Volume 


Table 3. Selected measurements (mm) and body mass (g) of the holotype of Myotis clydejonesi (TTU 109227), 
and of samples of M. cf. nigricans from Suriname and M. nigricans from Paraguay. Mean calculated for n > 3; 
n = sample size (adults only). See Methods for variable abbreviations and Appendix for localities of specimens. 


Variable 

TTU 109227, $ 

(Suriname) 

Holotype 

M. cf. nigricans 

(Suriname) 

Mean (Range), n 

M. nigricans 

(Paraguay) 

Mean (Range), n 

TL 

88 

- 

- 

Tail 

38 

- 

- 

Hind foot 

6 

- 

- 

Ear 

11 

- 

- 

Body mass 

4.3 

- 

- 

FA 

34.9 

35.0, 1 

32.9 (30.7-35.6), 54 

3ML 

33.6 

- 

31.1 (28.8-34.5), 54 

LDH 

7.7 

5.6, 1 

- 

lvh 

6.1 

4.5, 1 

- 

GLS 

14.3 

13.9(13.4-14.2), 3 

13.7(13.2-14.2), 53 

CCL 

12.8 

12.1 (11.7-12.3), 3 

12.0(11.7-12.5), 53 

CBL 

13.5 

12.7 (12.3-13.1), 3 

12.7(12.3-13.1), 52 

cil 

13.7 

12.9(12.5-13.2), 3 

12.9(12.5-13.3), 52 

bal 

12.3 

11.7(11.5-11.8), 3 

11.6(11.2-12.1), 52 

ZB 

8.8 

8.1, 8.3,2 

- 

MAB 

7.0 

6.9 (6.7-7.2), 3 

7.0 (6.8-7.3), 53 

BCB 

6.4 

6.6 (6.4-6.8), 3 

6.5 (6.2-6.9), 52 

IOB 

4.4 

4.6 (4.5-4.8), 3 

4.5 (4.0-4.8), 53 

POB 

3.4 

3.6 (3.6-3.7), 3 

3.6 (3.4-3.8), 53 

BAC 

3.5 

3.5 (3.4-3.7), 3 

3.5 (3.2-3.6), 41 

BAM 

5.7 

5.6 (5.5—5.7), 3 

5.3 (5.1-5.5), 41 

MTL 

5.3 

5.1 (5.0-5.2), 3 

5.0 (4.8-5.2), 52 

Ml-3 

3.0 

2.9 (2.8-3.0), 3 

2.9 (2.7-3.0), 53 

MAL 

10.2 

9.8 (9.5-10.1), 3 

9.6 (9.2-10.0), 53 

MAN 

5.6 

5.4 (5.3-5.5), 3 

5.3 (5.2-5.5), 53 


Moratelli et al.—New Myotis from Suriname 


59 


Figure 6. Map of part of South America illustrating the type locality (indicated by star) of Myotis clydejonesi at 
Raleigh Falls (04°43' N, 56°12' W), district of Sipaliwini, Suriname. 


the absence of a fringe of hairs along the trailing edge 
of the uropatagium; from M. oxyotus by the frontals 
smoothly inclined (not steeply sloping as in oxyotus ), 
and smaller external and cranial size (see Moratelli et 
al. 2013); from M. keaysi and M. riparius by the silky 
fur, and occipital region rounded; also, from M. keaysi 
by the dorsal fur on the uropatagium not reaching the 
knee, and the fur on the plagiopatagium along the 
body either absent or extremely sparse. Comparing 
M. clydejonesi with M. cf. nigricans from Suriname 
(using vouchers that were the sources of DNAfor our 
Cytb analysis), M. clydejonesi also can be distinguished 


by the flattened skull and elongated rostrum. These 
characteristics also distinguish M. clydejonesi from M. 
nigricans sensu stricto (Bolivia, Paraguay, E Brazil). 

Etymology.—Myotis clydejonesi honors Clyde 
Jones, in recognition of his outstanding contributions 
to mammalogy (see Jones 2005). Clyde was a mentor, 
colleague, supervisor, and friend; we find it particu¬ 
larly fitting that the type specimen of M. clydejonesi 
is housed in the Museum at Texas Tech University, the 
institution that holds a major part of his collections 
and legacy. 


60 


Clyde Jones Memorial Volume 


Discussion 


With the description of M. clydejonesi, 21 
formally described Neotropical species of Myotis 
currently are recognized (see Moratelli and Wilson 
2014). Husson (1962) recognized three species of 
Myotis from Suriname— M. albescens, M. nigricans, 
and M. surinamensis Husson, 1962. The latter he 
proposed as a replacement name for Vespertilio 
ferrugineus Temminck. However, according to 
Carter and Dolan (1978:73) and Davis and Gardner 
(2008:445), the type of V. ferrugineus does not represent 
a South American bat. Subsequently, Lim et al. (2005) 
reported three species ( albescens, nigricans, riparius), 
and M. clydejonesi now represents the fourth species 
for the country. Beyond the support for recognizing M. 
clydejonesi , our results also indicate that “M nigricans ” 
from Suriname possibly represents another undescribed 
species. 

Our findings also provide additional support for 
Larsen et al.’s (2012) hypothesis that M. nigricans 
should be restricted to southern South America. The 
species, as traditionally recognized, is polyphyletic. 
We suggest retaining the name “ nigricans ” for the 
haplogroup formed by Bolivian and Paraguayan 
samples because they are geographically closer to the 
type locality of the species in southeastern Brazil (see 
LaVal 1973; Moratelli et al. 2011) than is any of the 
remaining haplogroups. Based on our results, at least 
four geographic groups previously assigned to M. 


nigricans may require new names (Suriname, western 
Ecuador, eastern Peru, and Tobago; see Fig. 1). Our 
results require additional analyses, and we will return 
to collections for further morphological comparisons. 
However, based on the frequency with which we find 
single museum specimens that we cannot assign to 
any of the currently recognized species (e.g., TTU 
109227 when originally examined), we suspect that 
the diversity of Neotropical Myotis (~ 26 spp.) is still 
underestimated. 

The results of our phylogenetic and morphologi¬ 
cal analyses unquestionably ally M. clydejonesi with 
other species in the albescens group (sensu Moratelli et 
al. 2013). In northern South America and the Guiana 
Shield, this group comprises M. albescens, M. caucen- 
sis, M. clydejonesi, M. handleyi, M. nesopolus larensis, 
M. cf. nigricans, and M. oxyotus. Other Myotis found 
in the same region (M. keaysi, M. pilosatibialis, M. 
riparius, and M. simus) are in the ruber group. Finally, 
M. clydejonesi can be distinguished from all species in 
the Neotropical subclade (sensu Ruedi et al. 2013) by 
its combination of ventral and dorsal pelage color and 
the depressed braincase. Although not so flattened as 
in M. planiceps Baker, 1955, these two species share 
this trait (much more accentuated in planiceps ). The 
flattened braincase in these two species is an example 
of convergence; M. planiceps is a representative of 
the Nearctic subclade of Myotis (Haynie et al. 2016). 


Acknowledgments 


The following curators and collection staff 
provided access to specimens under their care: R. 
Timm (Biodiversity Institute, University of Kansas, 
Lawrence, USA); N. Simmons, E. Westwig (Ameri¬ 
can Museum of Natural History, New York, USA); 
J. Wible, S. McLaren (Carnegie Museum of Natural 
History, Pittsburgh, USA); C. Bernard (Centre for the 
Study of Biological Diversity, University of Guyana, 
Guyana); M. de Vivo, J. Gualda (Museu de Zoologia 
da Universidade de Sao Paulo, Brazil); R. Baker, H. 
Gamer (Museum of Texas Tech University, USA); M. 
Ruedi (Museum d’histoire naturelle, Geneva, Switzer¬ 
land); B. Lim (Royal Ontario Museum, Canada); K. 


Helgen, D. Lunde, and L. Gordon (Smithsonian’s Na¬ 
tional Museum of Natural History, USA). Support for 
Moratelli comes from the Brazilian National Council 
for Scientific and Technological Development / Science 
Without Borders Program (CNPq 202612/2012), and 
the Smithsonian Institution. Support for Fisher and 
Gardner comes from the USGS Ecosystems Mission 
Area. Support for E. Gutierrez comes from Univer¬ 
sidade de Brasilia, the Brazilian Coordination for the 
Improvement of Higher Education Personnel (CAPES), 
and the Smithsonian Institution. Any use of trade, prod¬ 
uct, or firm names is for descriptive purposes only and 
does not imply endorsement by the U.S. government. 


Moratelli et al.—New Myotis from Suriname 


61 


The holotype was obtained during fieldwork supported 
by funds from Outside Magazine granted to T. H. 
Genoways and funds from the Joyce Genoways Field 


Expedition Fund. We are grateful to T. H. Genoways, 
M. Bogan, and R. A. Van Den Bussche for helpfully 
reviewing previous drafts of the manuscript. 


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Addresses of authors: 

Ricardo Moratelli 

Fiocruz Mata Atlantica 

Fundagao Oswaldo Cruz 

Rio de Janeiro, RJ, CEP 22713-375 

Brazil 

rmoratelli@fiocruz. br 


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tion, University of Texas, Austin. 


Don E. Wilson 

Division of Mammals 
National Museum of Natural History 
Smithsonian Institution 
PO. Box37012 

Washington, DC 20013-7012, USA 
wilsond@si.edu 


Moratelli et al.—New Myotis from Suriname 


63 


Alfred L. Gardner 

USGS Patuxent Wildlife Research Center, Biological 
Survey Unit 

National Museum of Natural History 
P.O. Box 37012 

Washington, DC 20013-7012, USA 
gardnera@si.edu 

Robert D. Fisher 

USGS Patuxent Wildlife Research Center, Biological 
Survey Unit 

National Museum of Natural History 
PO. Box 37012 

Washington, DC 20013-7012, USA 
fisherr@si.edu 


Eliecer E. Gutierrez 

PPG Ecologia-PNPD 

Departamento de Zoologia 

Instituto de Ciencias Biologicas, Campus UnB 

Universidade de Brasilia 

Asa Norte, Brasilia, DF, CEP 70910-900 

Brazil 

gutierreze @ si. edu 


64 


Clyde Jones Memorial Volume 


Appendix 

Listed below are localities of specimens examined from northern South America, including northern Brazil, 
French Guiana, Guyana, Suriname, and Venezuela. They are preserved in the American Museum of Natural 
History (AMNH, New York, USA); Carnegie Museum of Natural History (CM, Pittsburgh, USA); Centre for 
the Study of Biological Diversity, University of Guyana (M, Georgetown, Guyana); Museum d’histoire naturelle 
(MHNG, Geneva, Switzerland); Museum of Texas Tech University (TTU, Lubbock, USA); Museu de Zoologia 
da Universidade de Sao Paulo (MZUSP, Sao Paulo, Brazil); National Museum of Natural History, Smithsonian 
Institution (USNM, Washington, DC, USA); and Royal Ontario Museum (ROM, Toronto, Canada). Localities 
are arranged alphabetically by species and major political unities. Specimens marked with asterisks were used 
in Table 2, and those with two asterisks also were used in the discriminant function analysis. 

Myotis albescens .—FRENCH GUIANA(l): Cayenne, Montsinery, Riviere des Cascades (MHNG 1990.017). 
GUIANA (18): Berbice, Dubulay (M 343); Essequibo, Chodikar River, 55 km SW of Gunn’s Strip (ROM 106655); 
Essequibo, Karanambo (ROM 97922); Potaro-Siparuni, Iwokrama Field Station, Iwokrama Forest (ROM 111997, 
112041, 112048, 112625); Potaro-Siparuni, Iwokrama Reserve, Pakatau Mountain, Siparuni River, 42 kmWNW 
ofKurupukari (ROM 107115); Potaro-Siparuni, Kabukalli Landing, Iwokrama Forest (ROM 111658); Rupumini, 
Kanukumi (M 177); Upper Takutu-Upper Essequibo, Dadanawa Ranch Headquarters (ROM 31892, 31903); 
Upper Takutu-Upper Essequibo, Dadanawa, Kuitaro River, Mountain on Right Bank (ROM 33002, 33003, 
33004, 33005, 33006, 33007). SURINAME (6): Nickerie, Grassalco (CM 63922, 63923, 63924, 63925, 63926); 
Saramacca, Raleigh Falls (CM 63928). VENEZUELA (3 9): Amazonas, Belen, 56 kmNNW of Esmeralda, Rio 
Cunucunuma (USNM 405790,405794,405796); Amazonas, Belen, 56 kmNNW of Esmeralda, Cano Essa (USNM 
405792); Amazonas, Capibara, 106 km SW of Esmeralda, Brazo Casiquiare (USNM 409392, 409395, 416579); 
Amazonas, Cerro Neblina Base Camp (USNM 560807, 560808); Amazonas, Paria, 25 km S of Puerto Ayacucho 
(USNM 409416, 409420, 409422, 409425); Amazonas, Rio Mavaca, 108 km SSE Esmeralda (USNM 405798); 
Amazonas, San Juan, 163 km ESE Puerto Ayacucho, Rio Manapiare (USNM 409403, 409404, 409406-409408, 
409410-409415, 409454, 416581); Amazonas, San Juan, Rio Manapiare (USNM 416580, 416582); Apure, Rio 
Cinaruco, 38 km NNW Puerto Paez (USNM 373909, 373913-373917, 374008); Apure, Nulita, 29 km SSW 
Santo Domingo, Selvas de San Camilo (USNM 441714-441716); Bolivar, Rio Supamo, 50 km SE El Manteco 
(USNM 387693); Miranda, 7 km E Rio Chico, near Puerto Tuy (USNM 387697-387701, 387703); Miranda, 10 
km SE Rio Chico, near Tacariquade La Laguna (USNM 387702); Trujillo, Valera, 23 kmNW Valera, Rio Motatan 
(USNM 370933); Zulia, El Rosario, 42 km NW Encontrados (USNM 441718). 

Myotis clydejonesi .—SURINAME (1): Sipaliwini, Raleigh Falls (TTU 109227** [holotype]). 

Myotis handleyi .—VENEZUELA (27): Aragua, Rancho Grande Biological Station, 13 kmNW Maracay 
(USNM 517503, 562923, 562924, 562925, 562926, 562927, 562928, 562929, 562930, 562931, 562932, 562933, 
562934,562935,562936,562937); Distrito Federal, Pico Avila, 5 kmNE Caracas, near Hotel Humboldt (USNM 
370932 [holotype]); Distrito Federal, Pico Avila, 5 kmNE Caracas, near Hotel Humboldt (USNM 370891 [para- 
type]); Miranda, Curupao, 5 kmNWGuarenas (USNM 387723); Monagas, 3 kmNWCaripe, near SanAgustin 
(USNM 409391, 409429, 409430, 409431, 409433, 409435, 409437, 409438). 

Myotis keaysi .—VENEZUELA (45): Aragua, Rancho Grande Biological Station, 13 km NW Maracay 
(USNM 370893-370895,370898-370902,370911-370913,370915-370922,370924,370926,370929); Aragua, 
Rancho Grande Biological Station, 13 km NW Maracay (USNM 370927, 370928, 370930, 370931); Aragua, 
Pico Guayamayo, 13 kmNW Maracay (USNM 521564); Aragua, Rancho Grande, Portachuelo (USNM 562920, 
563005, 563006); Aragua, Rancho Grande (USNM 562921); Bolivar, Gran Sabana (USNM 130625, 130626); 
Carabobo, Montalban, 4 km NW Montalban, La Copa (USNM 441741, 441742); Distrito Federal, Los Venados, 


Moratelli et al.—New Myotis from Suriname 


65 


4 km NW Caracas (USNM 370889); Distrito Federal, Pico Avila, 5 km NNE Caracas, near Hotel Humboldt 
(USNM 370890); Distrito Federal, junction Puerto Cruz Highway and Colonia Tovar Highway, 0.5 km W (USNM 
562984); Guarico, Hacienda El Vira, 10 km NE Altagracia (USNM 387707); Miranda, San Andres, 16 km SE 
Caracas (USNM 373920); Miranda, Curupao, 5 kmNWGuarenas (USNM 387714-387716, 387718); Monagas, 
Caripe (USNM 534265). 

Myotis nesopolus .—CURASAO (1): Punda area, Willemstad (USNM 101849 [holotype of M. nesopolus]). 
VENEZUELA (9); Falcon, Capatarida (USNM 441710, 441735-441737, 441740); Falcon, 6 km SW Capatari- 
da (USNM 441711); Falcon, Capatarida (USNM 441728); Lara, Rio Tucuyo (AMNH 130709 [holotype of M. 
larensis]); Zulia, Near Cojoro, 35 km NNE Paraguaipoa (USNM 441721). 

Myotis nigricans .—PARAGUAY (54): Presidente Hayes, 227 km NW Villa Hayes by road (MVZ 144707*, 
144708*, 144710*, 144711*, 144713*, 144714*, 144715*, 144716*, 144717*, 144719*, 144720*, 144722*, 

144726*, 144727*, 144728*, 144729*, 144730*, 144731*, 144732*, 144735*, 144738*, 144739*, 144741*, 

144743*, 144744*, 144746*, 144747*, 144748*, 144749*, 144750*, 144752*, 144753*, 144755*, 144756*, 

144757*, 144761*, 144762*, 144763*, 144764*, 144766*, 144767*, 144768, 144769, 144770**, 144771**, 

144772**, 144773**, 144774**, 144775**, 144776**, 144777**, 144778**, 144779*,144780*). 

Myotis cf. nigricans .—FRENCH GUIANA (7): (MHNG 1983.75, 1983.76, 1983.77, 1983.79, 1984.03, 
1984.05, 1990.54). GUYANA (35): Cuyuni-Mazaruni, Paruima (ROM 108263); Demerara-Mahaica, Ceiba 
Biological Center (ROM, 113797,112532, 112572, 112665); Upper Demerara-Berbice, Dubulay Ranch (USNM 
582351, USNM 582352); Upper Demerara-Berbice, Tropenbos, 20 km SSE of Mabura Hill (ROM 103479, 
103483); Upper Takutu-Upper Essequibo, Achimeriwau River, Mabi Wau, Near Achamere Wau (ROM 34042, 
34043, 34044, 34045, 34046, 34047, 34048, 34049); Upper Takutu-Upper Essequibo, Chipirari Wau Mouth, 15 
mi E of Dadanawa (ROM 34020); Upper Takutu-Upper Essequibo, Courchiwin Mountain, 10 mi E of Dadanawa 
(ROM 32890, 32892, 32893, 32894, 32896, 32897, 32900); Upper Takutu-Upper Essequibo, Essequibo River, 
7 km S of Gunn’s Strip (ROM 106738); Upper Takutu-Upper Essequibo, Gunn’s Strip (ROM 106772); Upper 
Takutu-Upper Essequibo, Karanambo (ROM 97931); Upper Takutu-Upper Essequibo, Komawariwau River, 
Comiwari Wau Mouth, 15 mi E of Dadanawa (ROM 34023, 34027, 34035, 34036); Upper Takutu-Upper Esse¬ 
quibo, Kuma River, 5 mi E, 5.5 mi S of Lethem, Kanuku Mountain (ROM 97827, 97828, 97879). SURINAME 
(3): Para, Zanderij (CM 63933**, 69053**, 77699**). VENEZUELA(64): Amazonas, Boca Mavaca, 84 km SE 
Esmeralda, 7 km up Rio Mavaca (USNM 405801); Amazonas, Paria, 25 km S Puerto Ayacucho (USNM 409424, 
409455); Apure, Nulita, 29 km SW Santo Domingo, Selvas de San Camilo (USNM 441722); Aragua, 3 km S 
Ocumare de La Costa (USNM 517504, 517505); Bolivar, Maripa (AMNH 17069 [holotype of M. maripensis ]); 
Carabobo, 10 kmNWUrama, El Central (USNM 373921, 373922, 373923, 373924, 373925, 373926, 373927, 
373928,373929,373930,373931,373932,373933,373934,373935,373936,373937-373941,373942,373943, 
373944,373945,373946,373947,373948,373949,373950,373951-373959,373989-374004); Carabobo, 6 km 
N Urama (USNM 374012); Trujillo, 11 km NW Urama, El Central (USNM 387708). 

Myotis oxyotus .—VENEZUEL A (9): Amazonas, Cerro Duida, Cano Culebra, 50 km NW Esmeralda (USNM 
405799); Amazonas, Cerro Neblina, Camp VII (USNM 560809-560811); Bolivar, Km. 125, 85 km SE El Dorado 
(USNM 3 87712); Bolivar, El Pauji, 21 km NE Icabaru, El Pauji (USNM 441750); Distrito Federal, Alto No Leon, 
33 km SW Caracas (USNM 409427); Merida, La Mucuy, 4 km E Tabay (USNM 373919, 387705). 

Myotis riparius .—FRENCH GUIANA(2): Paracou, near Sinnamary (AMNH 266376,268591). GUYANA 
(6): Barima-Waini, North West District (USNM 568021); Potaro-Siparuni, Iwokrama Field Station, Iwokrama 
Forest (ROM 112049); Potaro-Siparuni, Iwokrama Reserve, Burro Burro River, 25 km WNW of Kurupukari 
(ROM 107278,114620); Potaro-Siparuni, Mount Ayanganna, First Plateau Camp (ROM 114688,114689); Upper 


66 


Clyde Jones Memorial Volume 


Takutu-Upper Essequibo, Gunn’s Strip (ROM 106773). VENEZUELA (12): Amazonas, Boca Mavaca, 84 km 
SSE Esmeralda, 7 km up Rio Mavaca (USNM 405803, 405804); Amazonas, Capibara, 106 km SW Esmeral¬ 
da, Brazo Casiquiare (USNM 409457); Amazonas, ca. 2 km SE Cerro Neblina Base Camp (USNM 560625); 
Amazonas, Tamatama, Rio Orinoco (USNM 405806); Apure, Nulita, 29 km SW Santo Domingo, Selvas de San 
Camilo (USNM 416584, 441746, 441748); Aragua, Rancho Grande (USNM 562940); Barinas, 7 km NE Alta- 
mira (USNM 441743); Bolivar, Rio Supamo, 50 km SE El Manteco (USNM 387721); Bolivar, San Ignacio de 
Yhuruani (USNM 448544). 

Myotis simus .—BRAZIL (42): Amazonas, Borba (AMNH 91886-91892, 94224, 94225, 94227, 94230- 
94234); Amazonas, Itacoatiara (MZUSP 4372); Amazonas, Manaus (AMNH 79534, 91472-91478, 91500); 
Amazonas, Parintins (AMNH 92983, 93489-93497, 93922-93925); Amazonas, Rio Jurua (MZUSP 638, 1074). 


Non-volant Mammals of Ash Meadows National Wildlife Refuge, Nevada 

Richard W. Manning and Martin R. Heaney 


Abstract 

Ash Meadows National Wildlife Refuge is located within the Mojave Desert ecore- 
gion of southern Nevada. We conducted a mammal survey, across all habitat types, for 
two years using Sherman live-traps and pitfall traps. We comment on habitat associations 
(or preferences) and estimates of relative abundance for each species captured. Also, 
we comment on other large-bodied mammals of the Refuge. Three rodent species were 
especially abundant and accounted for approximately three-fourths of all live-trap and 
pitfall trap captures: MerrianTs Kangaroo Rat (. Dipodomys merriami ); Western Harvest 
Mouse (Reithrodontomys megalotis)\ and Cactus Deermouse (Peromyscus eremicus ). 

Key words: Ash Meadows National Wildlife Refuge, habitat preference, mam¬ 
mals, Nevada, rodent relative abundance 


Introduction 


During 2007-2009, a 2-year baseline wildlife 
inventory was conducted at Ash Meadows National 
Wildlife Refuge (hereafter, the Refuge or AMNWR) in 
Nye County, Nevada. This particular study is focused 
on the results of that survey that pertain to non-volant 
mammals on the Refuge. The main purposes of this 


study were to: 1) establish permanent sampling sites on 
the Refuge, including pitfall drift fence arrays (PDFAs) 
and Sherman® live-trap transects; and 2) collect base¬ 
line information on small ground-dwelling mammals 
on the Refuge. 


Study Site 


The AMNWR occurs entirely within the Mojave 
Desert ecoregion, which covers an area of approxi¬ 
mately 38,360 km 2 (14,811 mi 2 ) in southern Nevada 
(Fig. 1). Average annual precipitation is 65-190 mm 
(3-7 in). The flora is dominated by Creosotebush 
(Larrea tridentata). Allscale ( Atriplex polycarpa ), 
Brittlebush (. Encelia farinosa ), Desert Holly ( Atriplex 
hymenelytra ), White Burrobush ( Hymenoclea salsola ), 
and Joshua Tree ( Yucca brevifolia), as well as a number 
of common associated species. 

Ash Meadows National Wildlife Refuge is 
located in Nye County, Nevada, approximately 145 
km (90 mi) northwest of Las Vegas (Fig. 2). Several 
prominent physiographic features, often mentioned in 


text, are depicted in Figure 2. The Refuge was estab¬ 
lished during June of 1984, protecting nearly 9,300 ha 
(23,000 ac) of spring-fed wetlands and alkaline desert 
habitat. Ash Meadows Refuge supports at least 26 
endemic plants and animals, five of which are Feder¬ 
ally listed as Endangered. The Refuge is thought to 
harbor the largest concentration of terrestrial endemism 
in the continental United States (http://www.fws.gov/ 
desertcomplex/ashmeadows/). 

The Refuge consists of a complex mosaic of 
communities characterized by unique conditions re¬ 
lated to small-scale variations in floristic assemblages, 
topography, soil characteristics, drainage patterns, and 
other physical features of the landscape. Following the 


67 


68 


Clyde Jones Memorial Volume 


Toiyabe NF 
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Figure 1. Map depicting research area, Ash Meadows National Wildlife Refuge, within the state 
of Nevada. 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


69 


Figure 2. Schematic map of Ash Meadows showing major springs and points of interest. 
[Bostic, Joseph. Ash Meadows. 2000. miniQuest adventures for the part time explorer, http:// 
www.miniquest.com/blog/2000/11/4/ash-meadows. Accessed December 2015], 


70 


Clyde Jones Memorial Volume 


completion of vegetation mapping by the BIO-WEST 
botanical team, all habitat types were identified based 
on vegetation characteristics. 

Habitat Types 

Alkali Meadow. —Alkali Meadows exist in 
areas with a shallow water table (1-2 m) throughout 
the growing season. This community most often is 
located in a valley depression or at the bottom of al¬ 
luvial fans throughout arid deserts of the Southwest. 
The typical elevation range of Alkali Meadows at Ash 
MeadowsNWRis 1,070-2,130m(CNPS2009). Soils 
consist of alkaline clays and silts that often produce a 
salt ‘crust’ on the surface (Jones & Stokes Associates 
1993; UCSB 2009). Vegetation is low growing and 
consists of perennial grasses and sedges such as Alkali 
Sacaton ( Sporobolus airoides ), Saltgrass ( Distichlis 
spicata ), Beardless wildrye (Leymus triticoides ), and 
Arctic Rush ( Juncus arcticus ). Common shrubs within 
Alkali Meadows include Rubber Rabbitbrush (Ericam- 
eria nauseosa ) and Big Saltbush (Atriplex lentiformis ) 
(CNPS 2009). 

The most concentrated alkali communities 
occur in Carson Slough, the Crystal Reservoir and 
lower Crystal Marsh areas, and west of Big Spring. 
Throughout the Refuge, hydrology varies from satu¬ 
rated in the winter months to saturated year-around. Al¬ 
kali Meadow is the transitional habitat between Alkali 
Shrub-Scrub communities and the wetter habitat types 
such as Wet Meadow, Alkali Sink, Alkali Seep, and 
sometimes Emergent Marsh communities. Although 
Alkali Meadows may share many species in common 
with bordering communities, they are unique in hydrol¬ 
ogy, species densities, and species composition. Alkali 
Meadows differ from Alkali Seeps in that they have a 
much higher percentage of vegetation cover, and they 
differ from Wet Meadows in that they rarely, or for 
only a short time, have surface water. A salt crust is 
associated with Alkali Meadow habitats and appears 
to be more pronounced in depressions and areas with 
drastic annual hydrologic changes. More Black-tailed 
Jackrabbit ( Lepus califomicus) activity was observed in 
Alkali Meadow habitat than in any other habitat within 
the Refuge. Vegetation is moderate to dense with usu¬ 
ally more than 50% cover of herbaceous species. The 
two most common plant species here are Alkali Sacaton 


and Saltgrass. In some cases, both form monotypic 
stands. Other common plants not listed above but 
associated with Alkali Meadow on the Refuge are 
Sandburg Bluegrass (Poa secunda ), Mojave Thistle 
(Cirsium mojavense), Copperweed ( Oxytenia acerosa ), 
Whiteflower Rabbitbrush ( Chrysothamnus albidus ), 
and Mojave Seablite ( Suaeda moquinii). 

Alkali Playa. —Alkali Playas occur throughout 
the arid Southwest in the lowest elevations of desert 
basins where the topography is level to concave, and 
barren to sparsely vegetated (<10% cover) (Nature- 
Serve 2009). Playas form during periods of intermittent 
flooding and evaporation, which occur in high-ground¬ 
water years or after flash flood events (NatureServe 
2009; USGS 2009). Because of water evaporation, 
some playas maintain varying amounts of surface salt 
crust that limits the types of vegetation that can grow 
there. Soils have a characteristic clay layer or hard pan, 
which limits water drainage (NatureServe 2009). Plants 
common to Alkali Playas include spikerush (Eleocharis 
sp.), Iodinebush (. Allenrolfea occidentals), Mojave 
Seablite, Saltgrass, Alkali Sacaton, and Atriplex spe¬ 
cies (NatureServe 2009). Polygonal surface cracking 
is a common feature in Alkali Playas (USGS 2009). 

This habitat covers the least amount of area on the 
Refuge. It occurs in only one location, directly below 
Crystal Reservoir Dam. It appears that playa formation 
may be caused in part by seepage from Crystal Reser¬ 
voir Dam, as well as groundwater and seasonal rains. A 
salt crust with moderate to low thickness is associated 
with the Alkali Playa, as are scattered surface gravels. 
Vegetation density is low; the most common species 
are scattered sparsely throughout this habitat and con¬ 
centrated on its margins. These species are Shadscale 
(Atriplex confertifolia ), Arctic Rush, Saltgrass, Mojave 
Thistle, Whiteflower Rabbitbrush, and Alkali Sacaton. 

Alkali Seep. —Alkali Seep communities occur in 
unique areas across Southwestern deserts. The water 
table must be at or near the surface throughout the year 
for an Alkali Seep to form. Soils are slightly to heavily 
alkaline and do not allow ponding water during rain 
events (Nuzum2005; UCSB 2009). Alkali Seeps often 
are found bordering or within larger Alkali Meadow 
communities (NNHP 2009; UCSB 2009). Alkali 
Seeps occur on flats, sloping terrain, and drainages. 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


71 


Vegetation cover usually is low (Nuzum 2005; UCSB 
2009). Some common species in this habitat are Vel¬ 
vet Ash ( Fraxinus velutina ), Mesquite ( Prosopis sp.), 
Saltgrass, Shadscale, Arctic Rush, and Mojave Thistle 
(NNHP2009). 

This habitat occurs at low to mid elevations where 
the water table is near the surface. A moderate to high 
thickness of evaporated salt crust surface exists, which 
limits vegetative cover in comparison with neighboring 
Alkali Meadows. Soils are clay and saturated for most 
of the year. Small pockets of Alkali Seep occur within 
Alkali Meadow communities and near spring channels. 
Alkali Seeps may also transition into Wet Meadow and 
Emergent Marsh habitats. Alkali Seeps often form at 
the bottom of topographic breaks. Common species 
found in this habitat on the Refuge include those listed 
above, as well as Desert Polygala ( Polygala acantho- 
clada ), Whiteflower Rabbitbrush, Copperweed, and 
Alkali Sacaton. 

Alkali Shrub-scrub. —Alkali Shrub-scrub is a 
common habitat type found throughout the Mojave 
and Sonoran Deserts (Brown 1994), as well as in 
small stands in the Colorado and Great Basin Deserts 
(Rowlands 1988). It occurs from below sea level to 
1,800-m elevations and has deep soils high in silt 
and clay content that hold more water than soils of 
Creosote Shrublands. Alkali Shrub-scrub is described 
as occurring in two phases, a xerophytic phase and a 
halophytic phase. The distribution of each phase is 
influenced by groundwater availability, which in turn 
is based on topography and climate. The xerophytic 
phase is located at higher elevations and is composed 
of xeric shrubs, subshrubs, and few forbs or grasses. 
The halophytic phase occurs at lower elevations and 
often borders Alkali Play a, Alkali Sink, and Alkali 
Seep habitats. Because groundwater is more avail¬ 
able, a higher variety of plants are found in this phase, 
although they must be able to tolerate higher levels 
of alkalinity (Brown 1994). In both phases the plant 
community is dominated by plants in the goosefoot 
(Chenopodiaceae) family. The xerophytic phase is 
mostly dominated by Atriplex sp. The most common 
species are Four-wing Saltbush (Atriplex canescens ), 
Allscale, Desert Holly, and Shadscale (Rowlands 1988; 
CNPS 2009). In the halophytic phase the dominant 
species are still Chenopods but include more varieties 


such as Suaeda sp., Nitrophila sp., and Sarcobatus sp., 
as well as the introduction of grasses such as Alkali 
Sacaton and Saltgrass (Rowlands 1988; Brown 1994). 

The most common habitat type within the Ref¬ 
uge is Alkali Shrub-scrub. It occurs throughout the 
mid-elevations. It is found throughout the central and 
southern portions of the Refuge with both xerophytic 
and halophytic types. The halophytic phase is the most 
common on the Refuge and occurs at lower to mid 
elevations, often bordering washes, Alkali Meadows, 
Alkali Seeps, Alkali Sinks, or riparian areas. The most 
common dominant species in the halophytic phase are 
Alkali Goldenbush (Isocoma acradenia ), Shadscale, 
Rubber Rabbitbrush, Mojave Seablite, Mesquite 
(Prosopis pubescens ) and Honey Mesquite (Prosopis 
glandulosa). Soils are not well drained in either of the 
types because of high silt and clay content, but they 
are never saturated to the surface. Elevation change 
in the eastern section of the Refuge often is rapid and, 
therefore, the xerophytic type does not always develop; 
instead it transitions quickly to Salt Desert Scrub habitat 
or Creosote Shrubland. Other than the Atriplex spe¬ 
cies listed above, common species in the xerophytic 
phase are Desert Pepperweed (Lepidiumfremontii) and 
Threadleaf Snakeweed (Gutierrezia microcephala ). It 
is only in the driest of the xerophytic phase that some 
cactus species appear sparsely, such as Beavertail 
Pricklypear (Opuntia basilaris ) and Silver Cholla 
(Cylindropuntia echinocarpa ). The xerophytic phase 
of Alkali Shrub-scrub may have species in common 
with Salt Desert Scrub, but it has a higher vegetation 
density, occurs at a slightly lower elevation, does not 
have well-drained soils, and exhibits a more prominent 
forb and grass layer (see the Salt Desert Scrub habitat 
description). Although forb and grass species are 
more prominent in Alkali Shrub-scrub habitat than in 
Salt Desert Scrub, they are still a minor component. 
Common species include Buckwheat (Eriogonum sp.), 
Desert Globe-mallow (Sphaeralcea ambigua ), and 
Indian Rice Grass (Achnatherum hymenoides). 

Alkali Sink .—Alkali Sinks are found at low el¬ 
evations with high salinity and shallow water tables, 
typically less than three meters deep (Barbour et al. 
1977; CNPS 2009). Alkali Meadow and Alkali Shrub- 
scrub share many species in common with Alkali Sink 
habitats but differ in hydrology and soil. The surface of 


72 


Clyde Jones Memorial Volume 


Alkali Sinks is impermeable and, therefore, water pools 
there during rain events. As pools evaporate, a salt crust 
often develops (CNPS 2009). Mojave Seablite is the 
most common shrub in this habitat, and it can exist in 
large stands. Other common species include Grease- 
wood ( Sarcobatus vermiculatus ), Parry’s Saltbush 
(Atriplex parryi ), Tamarisk ( Tamarix sp.), Shadscale, 
Whiteflower Rabbitbrush, Alkali Sacaton, Saltgrass, 
Arrowweed, and Screwbean Mesquite. Allenrolfea 
species commonly occupy highly saline areas that 
other plants cannot tolerate (Barbour et al. 1977; CNPS 
2009). Alkali Sinks have many species in common with 
Alkali Playa; however, Alkali Playa have significantly 
less vegetation cover (<10%) (NatureServe 2009). 

Within the AMNWR, Alkali Sink habitats oc¬ 
cur at mid-elevation west and southwest of Peterson 
Reservoir, south of Crystal Marsh, and in scattered 
areas in the southern portion. Soils are clay and fine 
silt that are poorly drained with evidence of pooling 
water at the surface. There is no associated surface salt 
crust. Mojave Seablite is strongly dominant (> 50% 
coverage) in all Alkali Sink communities throughout 
the Refuge. Species found in Alkali Sink communities 
within the Refuge are less diverse than those found in 
regional habitats. Other than Mojave Seablite, common 
species associated with Alkali Sinks are Big Saltbush, 
Shadscale, Allscale, and Saltgrass. 

Ash. —Velvet Ash is a deciduous tree that pre¬ 
fers fine-textured soils and occurs in riparian areas 
throughout the Southwest (USDA2009). Although it 
is generally considered a riparian species, Velvet Ash is 
found both within and outside of the Refuge’s riparian 
habitats. For this reason, and because of the number 
of rare and endemic species found in Ash habitat, the 
Ash communities have been separated from Riparian 
Woodland (see Riparian Woodland habitat description) 
and assigned a separate habitat type. Ash habitat occurs 
throughout the low and mid-elevations of the Refuge, 
often within or bordering larger Alkali Meadow, Wet 
Meadow, Alkali Seep, and riparian habitats. The larg¬ 
est Ash communities within the Refuge occur on Col¬ 
lins Ranch, Mary Scott Spring, Scruggs Springs, and 
southeast of Crystal Reservoir. On the Refuge, Ash 
habitat most often occurs in small patches near springs 
and seeps, with one major exception, a community 
southeast of Crystal Reservoir that occurs within a 


larger Alkali Meadow community. This is the largest 
Ash population on the Refuge. Ash communities have 
a varying understory based on the surrounding habitat 
type in which they occur. Some common understory 
species are Alkali Sacaton, Big Saltbush, and Alkali 
Goldenbush. Other overstory trees that may be found 
within Ash habitat are Goodding’s Willow (Salix good- 
dingii ) and Screwbean Mesquite. 

Cottontop Dry Ridge. —Within the Mojave Des¬ 
ert, Cottontop Dry Ridge habitat is commonly described 
within Creosote Shrubland habitat based on similar 
species composition and environmental conditions (see 
Creosote Shrubland habitat description). The Cottontop 
Dry Ridge habitat classification is based on the 2006 
Ash Meadows Geomorphic and Biological Assessment 
final report (Otis Bay and Stevens Ecological Consult¬ 
ing 2006), which distinguishes Cottontop Dry Ridge 
as a habitat type within the Refuge based on a lower 
vegetative cover than in Creosote Shrubland, as well 
as the unique steep slopes and ridges that it occupies. 
Soil is poor and supports a low density of vegetation 
including Creosote Bush, White Bursage, Eriogonum 
species, Barrel Cactus ( Ferocactus cylindraceus ), and 
Silver Cholla. Cottontop Dry Ridge habitat occupies 
the highest elevations of the Refuge, the high slopes 
and ridges of the Specter Range bordering the eastern 
boundary of the Refuge. These Cambrian limestone 
and dolomite ridges rise steeply from the valley floor 
and transition quickly from Creosote Shrubland or Salt 
Desert Scrub into Cottontop Dry Ridge habitat (Otis 
Bay and Stevens Ecological Consulting 2006). 

Creosote Shrubland. —Creosote Shrubland is the 
single most dominant plant community found in the 
Mojave Desert (Brooks et al. 2007). It occurs at less 
than 1,220 m in elevation above the “saltbush zone,” 
which is dominated by Atriplex species, and below the 
“Blackbrush Zone,” which is dominated by Blackbrush 
(Coleogyne ramosissima ) (Brown 1994). Creosote 
Shrubland often occupies broad valleys, plains, low 
hills, and lower bajadas. The characteristic soil is well- 
drained sand with large surface gravels often forming 
desert pavement (NatureServe 2009). Creosotebush 
is the dominant shrub and most often associated with 
white bursage; however, other shrubs may occupy and 
dominate the community at varying elevations. Some 
common, co-dominant species are Shadscale, Desert 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


73 


Holly, Ephedra species, Encelia species, and Desert- 
thorn (NatureServe 2009). These communities are 
highly prone to fire damage (Brooks et al. 2007). 

The highest elevations on the Refuge are occupied 
by Creosote Shrubland habitat. It is distributed on the 
east of the Refuge and is similar to Creosote Shrubland 
found throughout the Southwest. Creosote Shrubland 
habitat differs from Salt Desert Scrub in that it occurs 
at slightly higher elevation. Although they may have 
some species in common, Creosote Shrubland habitat 
is dominated by Creosotebush and White Bursage, with 
a minor component of Shadscale, Desert Holly, Button 
Brittlebush (Encelia frutescens), White Ratany (Kra¬ 
mer ia grayi), and Rusty Molly (Bassia californica). 
Soil is well drained with dark cobbles on the surface 
that often form desert pavement. The herbaceous layer 
is sparse but may include Desert Trumpet (Eriogonum 
inflatum ), Rigid Spineflower (Chorizanthe rigida ), and 
winter annuals such as Phacelia species. 

Dunes. —Stabilized Dunes and Coppice Dunes 
are found in warm, semi-arid regions of the Southwest. 
Soils commonly associated with dunes are quartz or 
gypsum very fine- to medium-grained sands (Nature- 
Serve 2009; USACE 2009). Within a given area, dune 
size is mostly uniform and can range from 0.5 to 3 m 
high and 1 to 15 m wide. Dunes form in areas where 
vegetation acts as nets, catching sands that build up in 
mounds. Vegetation that stabilizes dunes must be able 
to tolerate having its branches and roots continually 
covered with sand. In the Southwest the most com¬ 
mon stabilizing species is Honey Mesquite (Rango et 
al. 2000). Other species found in dune communities 
are Mormon Tea (Ephedra sp.), Four-wing Saltbush, 
Acacia species, Tamarix species, White Bursage, Desert 
Sand Verbena (Abronia villosa ), Sand Sagebrush (Ar¬ 
temisia filifolia ), Dune Buckwheat (Eriogonum deser- 
ticola), Sporobolus species, and Creosotebush (Rango 
et al. 2000; NatureServe 2009; USACE 2009). 

Within the Refuge there are two types of dune; 
Mesquite Dunes and Shrub Dunes. Both are Coppice 
Dunes, each with different plants acting as the stabiliz¬ 
ing species. Mesquite Dunes are stabilized by honey 
mesquite and are the most common dunes found on 
the Refuge. Shrub Dunes are stabilized by four-wing 
saltbush and are less abundant than Mesquite Dunes. 
Heavy winds form dunes at mid-elevations across 


the Refuge with a concentration on the western side. 
Soils are well drained and sandy. Dune communities, 
especially Mesquite Dunes, are highly productive ar¬ 
eas where much animal activity and cultural resources 
(lithic scatter) have been observed. The largest acreage 
of Mesquite Dunes is found west of Horseshoe Marsh 
and west of Cold Spring. Because of the high winds 
and xeric conditions on the Refuge, dune-forming 
honey mesquite are low and shrub-like in most areas. 
Elsewhere in the Southwest, nondune-forming Honey 
Mesquite are mostly upright with a clear central stem. 
Common species within a Mesquite Dune community 
are Honey Mesquite, Atriplex species, Alkali Gold- 
enbush, Rubber Rabbitbrush, Mojave Seablite, and 
Alkali Sacaton. Habitat interlaced within dunes most 
commonly resembles Alkali Shrub-scrub. Dune com¬ 
munities support a variety of winter annuals including 
Eriogonum species and Booth’s Primrose (Camissonia 
boothii). The largest area of Shrub Dunes is found at the 
western edge of Horseshoe and Crystal marshes, as well 
as west and south of Peterson Reservoir. Vegetation 
usually is less dense than in Mesquite Dunes, and these 
communities support a lower diversity of plants. Com¬ 
mon species include Four-wing Saltbush, Thurber’s 
Sandpaper Plant (Petalonyx thurberi ), shadscale, and 
Alkali Goldenbush. 

Emergent Marsh. —Emergent Marshes are dis¬ 
tributed widely throughout all elevations in the arid 
West but are most concentrated below 2,270 m (Kramer 
1988; NatureServe 2009). Emergent Marshes occur on 
all slopes but most often occur in depressions on the 
landscape or across level or rolling terrain. Frequent to 
continual inundation of Emergent Marshes, with water 
1-2 m deep, results in hydric silt and clay soils that may 
display gleyed coloring, high amounts of organic mat¬ 
ter, and/or redoximorphic features (NatureServe 2009). 
This vegetation community is dominated by perennial 
hydrophytic herbaceous genera such as Schoenoplectus, 
Typha, Juncus, and Phalaris. Species differ by region 
(NatureServe 2009). Emergent Marshes often are bor¬ 
dered by Wet Meadows marked by the transition from 
hydrophytic to mesophytic vegetation. Deep-water 
habitat may occur within an Emergent Marsh when the 
water level reaches more than two meters and emergent 
vegetation can no longer survive (Kramer 1988). The 
amount of open water within an Emergent Marsh will 
vary from season to season (Kramer 1988). 


74 


Clyde Jones Memorial Volume 


Habitat designated as Emergent Marsh within 
the AMNWR is most abundant at lower Crystal and 
Horseshoe marshes. Area of Emergent Marsh will 
vary from year to year based on annual rainfall and 
temperature. Crystal and Peterson reservoirs also 
support Emergent Marsh habitat. Carson Slough has a 
lattice of Emergent Marsh vegetation that is dominated 
by Common Reed ( Phragmites australis). Much of the 
common reed stands appeared to be drying out because 
of changing hydrology in the area. Other Emergent 
Marsh communities on the Refuge are associated with 
spring outflows including Bradford, Big, and Kings 
springs. Common species are Typha species, Chair- 
maker’s Bulrush ( Schoenoplectus americanus ), and 
Common Reed, with Arctic Rush, Beaked Spikerush 
(Eleocharis rostellata ), and Saltgrass bordering the 
edge of Emergent Marsh communities as they transi¬ 
tion to Wet Meadows. 

Mesquite Bosque. —Mesquite Bosque communi¬ 
ties are found along rivers in the Southwest, including, 
but not limited to, the Colorado River, Gila River, 
Santa Cruz River, and Rio Grande (NatureServe 2009). 
Although Mesquite Bosques are considered riparian 
habitat, they often occur somewhat distant from streams 
on sites with less reliable hydrology (e.g., alluvial ter¬ 
races, washes, and alkali sinks) but are dependent on 
the seasonal rise in groundwater (NatureServe 2009; 
UCSB 2009). Stands are distributed on toe slopes or 
valley bottoms at elevations below 1,100 m (Nature¬ 
Serve 2009). The canopy consists of Prosopis species, 
including Honey Mesquite, Velvet Mesquite ( Prosopis 
velutina), and Screwbean Mesquite. Common spe¬ 
cies occupying the shrub layer, when one exists, are 
Baccharis species and Coyote Willow ( Salix exigua). 
The herbaceous layer is generally open but may con¬ 
tain grass species and Atriplex species (NatureServe 
2009; UCSB 2009). Mesquite Bosques are important 
habitat for many mammals, birds, insects, and reptiles 
throughout the Southwest, providing shade and food 
resources (Plagens 2009). 

Within the Refuge, Mesquite Bosques are dis¬ 
tributed throughout low elevations with higher con¬ 
centrations occurring near springs, seeps, and washes. 
Because of the high groundwater table in the Refuge’s 
low elevations, Mesquite Bosques are not restricted to 
riparian areas. Areas near spring channels dominated 


by screwbean mesquite have been designated as Ri¬ 
parian Woodland habitat to highlight their proximity 
to water and distinguish them from mesquite stands 
not associated with a channel. All mesquite stands 
not directly adjacent to spring channels have been 
designated as Mesquite Bosque habitat. Screwbean 
Mesquite is the most common dominant species, with 
honey mesquite comprising a portion of the canopy in 
some areas. Other associated tree species are Tama¬ 
risk, Velvet Ash, and Fremont Cottonwood ( Populus 
fremontii). The most common understory species are 
Saltgrass, Big Saltbush, Golden Alkalibush, Alkali 
Sacaton, and Shadscale. Canopy height of this com¬ 
munity averages 3-6 m and provides important habitat 
for animals in the area. 

Nonnative/Weed. —Nonnative/Weed habitat is 
assigned to communities dominated by species that 
are not native to the Refuge. This excludes Tamarisk, 
which has been placed in its own habitat type because 
of the unique management challenge it poses for the 
Refuge. Transitioning Agricultural habitat also may 
be dominated by nonnative species; however, Transi¬ 
tioning Agriculture occurs only on historic agricultural 
fields, whereas Nonnative/Weed habitat occurs in other 
disturbed areas such as roadsides, historic restoration 
areas, and on former private property. Some Non¬ 
native/Weed habitat, such as Carson Slough, falls 
within historic agricultural fields but no longer shows 
evidence of agriculture in the field. Nonnative/Weed 
communities exist mostly in wet areas across the low 
to mid-elevations of the Refuge, including the west 
side of Carson Slough, Bradford Springs area, and 
south of Kings Spring. Dominant plants include Five- 
hook Bassia ( Bassia hyssopifolia ), Russian Knapweed 
(.Acroptilon repens ), Redstem Stork’s Bill ( Erodium 
cicutarium ), Common Sunflower (Helianthus annuus), 
and Spreading Alkaliweed. Russian Knapweed was 
being actively controlled during mapping. Habitat des¬ 
ignations represent community composition at time of 
data collection and may not reflect current conditions. 
Common Sunflower and Spreading Alkaliweed are 
native species that have been included because of their 
apparent preference and success on disturbed surfaces. 

Riparian Woodland and Shrubland. —Desert 
Riparian Woodlands and Shrublands are important 
habitats located along perennial streams and spring 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


75 


outflows throughout the Mojave and Sonoran deserts 
(Laundenslayer 1988). Riparian zones are dependent 
on seasonal flooding provided by the streams they 
occupy (NatureServe 2009). The soils range from 
rocky, sandy, well-drained soil to silty alluvial deposits 
(Laundenslayer 1988). Desert Riparian Woodlands and 
Shrublands occur at low elevations of less than 1,200 m 
in canyons or valley bottoms. Tree canopy commonly 
is dominated by Fremont Cottonwood, Goodding’s 
Willow, Velvet Ash, Prosopis species, and Tamarisk. 
The shrub layer, which may be an understory to the 
tree layer or form unique stands along stream chan¬ 
nels, often is composed of Big Saltbush, Narrowleaf 
Willow (, Salix exigua), arrowweed, or Mojave Seablite 
(Laundenslayer 1988; NatureServe 2009; USFS 2009). 
Desert Riparian areas average 7.5-24.5 mm (3.1-9.8 
in) precipitation a year, but communities may survive 
in much drier conditions if the water table is seasonally 
available (Laundenslayer 1988). 

Riparian Woodlands on the Refuge are associated 
with spring discharges and their surrounding areas. 
The Riparian Woodlands classification includes two 
types, those dominated by Prosopis species and those 
dominated by Salix and/or Populus species. Because 
Mesquite Bosque communities are found throughout 
the Refuge in non-riparian areas, communities domi¬ 
nated by Prosopis species in riparian areas were desig¬ 
nated as Riparian Woodland, and those dominated by 
Prosopis species in non-riparian areas were designated 
as Mesquite Bosque (see Mesquite Bosque habitat de¬ 
scription). Common tree species are the same as above 
with the exception of velvet ash, which has been placed 
in a unique habitat type (see Ash habitat description). 
The largest amount of Riparian Woodland occurs along 
the channel corridor of Kings Spring and other nearby 
springs, as well as in the Scruggs Springs area, Crystal 
Spring channel, and southwest of Bradford Springs. 
Canopy cover is dense with varying understory com¬ 
position. The most common understory plants are Big 
Saltbush and Alkali Sacaton. Riparian Woodland most 
often transitions into Riparian Shrubland or Alkali 
Shrub-Scrub communities. 

Riparian Shrublands are associated with spring 
outflow in the same areas as or independently of Ripar¬ 
ian Woodlands throughout the Refuge. They are either 
the understory of Riparian Woodlands or occur in inde¬ 


pendent stands. When the two habitats occur together, 
the Riparian Woodlands will be nearest to the spring 
channel and transition to Riparian Shrublands farther 
back from the spring channel. Riparian Shrublands 
in the vicinity of Big Spring occur independently of 
Riparian Woodlands. Common shrubs are as indicated 
above with the addition of Emory’s Baccharis ( Bac- 
charis emoryi), which is common in the Refuge both 
within and outside of riparian areas. 

Salt Desert Scrub. —Salt Desert Scrub, also 
known as Shadscale Scrub, occurs throughout the Mo¬ 
jave and Great Basin deserts on low slopes of alluvial 
fans. The soils generally are poorly drained and slightly 
alkaline. Plant density is low and canopy height ranges 
from 0.3 to 1 m. Salt Desert Scrub occurs below Creo¬ 
sote Shrubland, which is dominated by Creosotebush, 
in the Mojave Desert and below Sagebrush Scrub, 
which is dominated by Artemisia species, in the Great 
Basin Desert (Smith 2000). Some of the most common 
species associated with this community are Shadscale, 
Blackbrush, Budsage (. Artemisia spinescens ), Desert 
Alyssum (Lepidium fremontii), Four-wing Saltbush, 
Fremont’s Dalea (Psorothamnus fremontii), Threadleaf 
Snakeweed, Ephedra species, Spiny Hopsage, Spiny 
Menodora, and Winterfat. Salt Desert Scrub has many 
species in common with Alkali Shrub-Scrub but is 
found at slightly higher elevations and in more xeric 
conditions (Smith 2000). 

Salt Desert Scrub is one of the driest habitat types 
in the Refuge, second only to Creosote Shrubland. It 
occurs on the eastern edge of the Refuge on toe slopes, 
alluvial fans, and badlands, transitioning quickly from 
the xerophytic phase of Alkali Shrub-Scrub. Salt Desert 
Scrub also can be found southwest of Cold Spring on 
the westernmost boundary of the Refuge. The most 
notable difference between Salt Desert Scrub and 
Alkali Shrub-Scrub communities are that Salt Desert 
Scrub has well-drained soils, which often form desert 
pavement, along with low vegetation density, minimal 
forb and grass layer, and a slightly higher elevation. 
Common species within the Refuge are as above with 
the exception of Blackbrush, which was only observed 
once on the refuge. The herbaceous layer is sparse. 
The two most common species are Desert Trumpet 
and rigid Spineflower, which occur along with other 
winter annuals. 


76 


Clyde Jones Memorial Volume 


Tamarix.—Tamarix communities form dense 
(60-100% cover) stands along riparian corridors 
throughout the Southwest. Communities often are 
monotypic with few non -Tamarix species contributing 
to overall cover (Hart 2009; USFS 2009). Tamarix 
is most successful below sea level to 2,000 m but 
has been found at elevations up to 3,350 m (USFS 
2009). Tamarix has the ability to occupy non-riparian 
areas because of its extensive root system, which may 
reach deep groundwater otherwise unavailable to na¬ 
tive vegetation (Hart 1999). These communities are 
tolerant of a wide variety of soil conditions and can 
withstand high salt concentrations. Tamarix often 
occurs in previously disturbed areas. The two main 
Tamarix species widespread throughout the South¬ 
west are Small Flower Saltcedar (T. parviflora) and 
Saltcedar ( T. ramosissima ). Other species that may 
occur in Tamarix stands are Arrowweed, Fremont 
Cottonwood, Narrowleaf Willow, Goodding’s Willow, 
Prosopis species, and Big Saltbush. Annual grasses 
may occupy the understory (Hart 1999; USFS 2009). 
Tamarix is a non-native species that was introduced to 
the United States in the 1800s. It has since become one 
of the most common riparian species in the Southwest. 
Some of the effects of Tamarix communities include 
displacement of native species, increased soil salinity, 
increased water consumption, increased fire frequency 
due to the high amount of fuel load, and increased flood 
events (Hart 1999). 

Tamarix habitat within the Refuge ranges from a 
few individuals along washes to large stands composing 
100% of the canopy. Tamarix was found throughout 
the Refuge in varying soil conditions in proximity 
to open water, ephemeral washes, or areas of high 
groundwater. Carson Slough, southeast of Jackrabbit 
Spring, south of Kings Spring, and around Peterson 
and Crystal reservoirs had the most Tamarix. The 
period of data collection often did not correspond to 
Tamarix flowering time and, therefore, Tamarix was 
not identified to the species level. Throughout the 
mapping process the Refuge was actively controlling 
the presence of Tamarix. Habitat designations were 
made based on the dominant vegetation at the time 
of data collection, and they may or may not represent 
current conditions. In many areas where Tamarix has 
been removed, it was observed that Tamarix seedlings 
were resprouting or old stumps were greening from the 
base. Common species associated with Tamarix are 


Screwbean Mesquite, Fivehook Bassia, Emory’s Bac- 
charis, Shadscale, Saltgrass, and common sunflower. 
Continued control of Tamarix is needed throughout the 
Refuge, especially in active restoration areas such as 
Peterson and Crystal reservoirs. 

Transitioning Agriculture. —Transitioning Ag¬ 
riculture habitat was assigned when a community 
occurred on a historic agricultural field but did not 
yet resemble or function as any other habitat type. 
Transitioning Agriculture was scattered throughout the 
mid elevations of the Refuge in areas such as Bradford 
Springs, west of Point of Rocks Springs, southwest of 
Jackrabbit Spring, west of the Refuge office, and in the 
southern portion of Carson Slough. Most Transitioning 
Agriculture areas were at varying levels of succession, 
and many still had plow lines or planting rows visible on 
the ground and in aerial photographs. Vegetative cover 
is low profile and often has a nonnative component. 
Other historic agricultural fields within the Refuge 
that are not assigned as Transitioning Agriculture are 
further along in transition and resemble an appropriate 
habitat type. Composition varies based on surrounding 
communities and location on the Refuge. Some native 
species that appear to prefer (or are successful on) the 
disturbed surfaces of Transitioning Agriculture are 
Brownplume Wirelettuce ( Stephanomeria pauciflora ), 
Emory’s Baccharis, Big Saltbush, and Honey Mesquite. 
Nonnative species include Desert Indianwheat ( Plan - 
tago ovata ) and Redstem Stork’s Bill. As succession 
and restoration continue, these areas may become more 
naturalized and act as functioning habitat types. 

Wet Meadow. —Wet Meadows occur throughout 
the western United States and are characterized by a 
dense herbaceous layer (60-100% cover) and little to 
no tree or shrub layer (Ratliff 1988). The hydric soil 
associated with Wet Meadows is poorly to moderately 
drained with a texture of clay loams to fine sands 
(MTNHP 2009). The water table is at the surface for 
most of the growing season but can fall to as much as 
a meter below the surface during the dry season, espe¬ 
cially in the Southwest. Wet Meadows occupy seeps, 
alluvial terraces, stream benches, overflow channels, 
and areas near springs on level to slightly undulating 
surfaces; they also often transition to Emergent Marsh 
communities (Ratliff 1988; MTNHP 2009). Associ¬ 
ated species are Arctic Rush, Sedge ( Carex sp.), Tufted 
Hairgrass (. Deschampsia caespitosa ), Saltgrass, Bui- 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


77 


rush ( Schoenoplectus sp.), Eleocharis species, Foxtail 
Barley ( Hordeum jubatum ), and Alkali Sacaton (Ratliff 
1988; MTNHP 2009). 

Wet Meadows occur throughout low elevations 
of the AMNWR where the water table is at or near the 
surface during most of the season, in open meadows, 
and in drainages associated with spring outflow. Carson 
Slough, southwest of Crystal Reservoir, and west of 
Big Spring are the largest areas of Wet Meadow habitat 
within the Refuge. Wet Meadows often are the transi¬ 
tion between mesic Alkali Meadow and hydrophytic 
Emergent Marsh habitats. Where Wet Meadows occur 
in topographically low areas or drainages, they may be 
bordered by Alkali Shrub-Scrub or other, more mesic 


habitat types. Although Wet Meadows may share 
many species in common with Alkali Meadow, Wet 
Meadow soils are saturated to the surface for much 
of the growing season. Plant species associated with 
Wet Meadows must be able to tolerate inundation as 
well as short dry periods. The most common species 
associated with Wet Meadows within the Refuge are 
Eleocharis species, Juncus species (most commonly 
Arctic Rush), Sandburg Bluegrass, and Saltgrass, with 
scattered Chairmaker’s Bulrush and Sturdy Bulrush 
0 Schoenoplectus robustus ). Shrubs and trees mostly 
are absent from Wet Meadows with the occasional 
Emory’s Baccharis, Big Saltbush, Screwbean Mesquite, 
and Velvet Ash at the edges. 


Materials and Methods 


Pitfall drift fence arrays. —Pitfall drift fence ar¬ 
rays (PDFAs) were the primary method used to sample 
amphibians and reptiles on the Refuge and incidentally 
take small mammals. Bury and Com (1987), Corn and 
Bury (1990), and Corn (1994) suggest that PDFAs are 
an effective way to sample species richness and deter¬ 
mine cryptic and rare species. The PDFAs consisted 
of a central 6-gallon plastic bucket with the rim buried 
flush with the ground and three 10-m spans of galva¬ 
nized sheet metal flashing placed as vertical barriers 
radiating out (hereafter, “arms”). The configuration of 
PDFAs varied slightly with location, depending on the 
surrounding vegetation and topography; however, the 
main design was in the shape of a “Y,” with one arm 
extending due north and the remaining arms radiating 
out from the center with equal angles of approximately 
120 degrees (Fig. 3). In addition to the center, pitfall 
traps were placed in the middle and at the end of each 
10-m arm. All pitfall trap buckets had snap-on remov¬ 
able lids. Funnel traps were placed between the middle 
and end of each arm. Loose soil, sand, and debris were 
placed at the bottom of each pitfall to provide protec¬ 
tion for captured specimens. The number of PDFAs 
by habitat type are presented in Table 1. 

Waypoints (UTM NAD 83, Zone 11) were col¬ 
lected for the location of every PDFA(see PDFA local¬ 
ity map, Fig. 4) installed on the AMNWR and entered 
into the database. Each pitfall trap open for one night 
was considered one trap-night. 


Field efforts were conducted over two years dur¬ 
ing three seasons: spring (6-11 April 2008 and 5-10 
April 2009); summer (1-6 June 2008 and 31 May-4 
June 2009); and fall (19-24 October 2008 and 18-23 
October 2009). Prior to starting the second-year field 
efforts, a PDFA repair trip was conducted 25-27 Febru¬ 
ary 2009. Repairs consisted of replacing pitfall bucket 
lids, reattaching flashing to rebar or wooden lathes, and 
replacing galvanized flashing that was fatigue-cracked 
by strong winds. 

Sherman live-trapping. —Sherman live-traps 
were the primary trap used to survey small mammals 
on the Refuge (see Jones et al. 1996; Wemmer et al. 
1996). Trap-nights are defined as one trap set for one 
night. Two field teams each deployed approximately 
180-200 traps per night along transects located in vari¬ 
ous habitat on the Refuge. Traps were retrieved the 
following morning and deployed at another location. 
Traplines consisted of approximately 30-40 traps per 
line with individual trap stations positioned approxi¬ 
mately 10 m (33 ft) apart. One or two traps were set at 
each station and baited with oatmeal. In many cases, 
trapline starting points were associated with a terminal 
PDFA pitfall bucket. Waypoints were collected for each 
trapline end point. 


78 


Clyde Jones Memorial Volume 


& - Funnel trap 
O - 6-ga!lon pitfall bucket 


Drift Fence Array Plan View 


Drift Fence Array Profile View 

Figure 3. Schematic of the pitfall drift fence arrays (PDFAs) installed at Ash Meadows National 
Wildlife Refuge, Nevada. 


Table 1. The number of pitfall drift fence arrays (PDFAs) in 
each habitat type at Ash Meadows National Refuge, Nevada, 
2008-2009. 


Habitat Type 

Number of PDFAs 

Alkali Shmb Scrub 

11 

Creosote Shrubland 

5 

Dune (Mesquite) 

3 

Alkali Meadow 

2 

Transitioning Agriculture 

2 

Alkali Sink 

1 

Ash 

1 

Dune (Shmb) 

1 

Mesquite Bosque 

1 

Riparian Shrubland 

1 

Riparian Woodland 

1 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


79 


1—'—1— 1 —1 > 

0 1 Kilometer A 


1 T T ’ 1 ^ 

0 1 Mile 


Job: 

Date: 

lulj'21,2009 

1 inch - fnt 

DfiTOtTl ht“ 

T. SchfttiiMf 

Fir. 


Figure 4. Location of pitfall drift fence arrays (PDFAs) on Ash Meadows National Wildlife Refuge. These 
locales served as the foci of Sherman live-trap transect lines. 


80 


Clyde Jones Memorial Volume 


All live specimens captured were identified to 
species and sex, released on-site, and the data were 
recorded. Ambient environmental data also were col¬ 
lected at each sampling location, including temperature, 
wind speed, wind direction (using handheld Kestrel 
anemometer), percent cloud cover, and precipitation. 

Game cameras. —Four motion-activated Moult¬ 
rie i40 digital game cameras were used to inventory 
additional medium to large mammals. Cameras were 
rotated among habitats each season. Cameras were 
mounted on steel T-posts and left in habitat types 
between seasonal sampling events. Each camera sta¬ 
tion was treated with scent attractants to increase the 
likelihood of mammal observation. Mammal species 
observed with the game cameras were recorded. Am¬ 
bient environmental data were collected by the game 
camera, which included start and stop times, date, and 
temperature. 

Incidental observations. —A number of qualita¬ 
tive efforts were employed in addition to the Sherman 
live-trap sampling efforts. All small mammals captured 
in PDFAs were recorded. Small mammal incidental 
observations were recorded and waypoints collected 
at each encounter. Road cruises were conducted near 
dawn and dusk to document mammal species. A GPS 
waypoint was collected for each mammal encounter. 


Tissue samples. —Prior to release of live speci¬ 
mens, tissue samples (ear clips) were collected using a 2 
mm (0.1 in) diameter punch. No more than 30 samples 
per species were collected. Tissues were placed in a 
Nunc® tube and stored in 95% ETOH. Any trap-dead 
mammal was prepared as a voucher specimen (usu¬ 
ally skin and skull); skeletal muscle, heart, and liver 
tissue also were collected. All tissue samples were 
deposited at the Natural Science Research Laboratory 
of the Museum of Texas Tech University (TTU) in 
Lubbock, Texas. 

Analysis of relative abundance. —Quantitative 
population and community metrics for small mammals 
were calculated using only Sherman live-trap data. 
Observed relative abundance is defined as the number 
of individuals captured per trap-night. 

Known collection specimens. —We know of 
mammal material in three museum collections from 
Ash Meadows: United States Museum of Natural 
History (NMNH), Smithsonian Institution; Museum 
of Vertebrate Zoology at Berkeley (MVZ); and the 
Natural Science Research Laboratory of the Museum 
of Texas Tech University (TTU). These specimens are 
listed after the individual species accounts. 


Results 


During our small mammal trapping survey (rep¬ 
resenting 10,910 trap-nights) the overall trap success 
rate was 10.4% (i.e., 10.4 animals per 100 trap-nights) 
across all habitat types. A total of 1,130 individuals 
representing 15 species were captured in Sherman traps 
(Table 2). Three species accounted for 72.3% of all 
captures. MerrianTs Kangaroo Rat ( Dipodomys mer- 
riami ) was the most commonly captured small mammal 
(.n = 276, or 24.4% of all captures); Western Harvest 
Mouse (Reithrodontomys megalotis) was the second 
most numerous small mammal taken (n = 275, or 24.3% 
of all captures); and the Cactus Deermouse ( Pero- 
myscus eremicus) ranked as the third most commonly 
trapped rodent (n = 267, or 23.6% of all captures). 


A total of225 individuals representing 14 species 
(1 soricomorph, 1 lagomorph, and 12 rodents) were 
captured in pitfall traps (Table 3). The three most fre¬ 
quently captured species were the Cactus Deermouse (n 
= 112, or 49.8% of captures); Western Harvest Mouse 
(n = 31, or 13.8% of captures); and Southern Grasshop¬ 
per Mouse ( Onychomys torridus) (n = 25, or 11.1% of 
captures). These three species accounted for 74.7% of 
all PDFA small mammal captures. 

Accounts of Species 

Common and scientific names follow Bradley 
et al. (2014) and Mantooth and Riddle (2005) unless 
otherwise noted. 


Table 2. Number of small mammals captured in Sherman traps (along with total trap-nights) by habitat type, at Ash Meadows National Wildlife Refuge, 
2008-2009. 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


81 


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Table 3. Number of small mammals captured in pitfall drift fence arrays (along with total trap-nights) by habitat type, at Ash Meadows National 
Wildlife Refuge, 2008-2009. 


Clyde Jones Memorial Volume 


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Sylvilagus audubonii 2 2 

Thomomys bottae 2 1 14 

Total captured 3 0 53 3 20 0 22 20 20 0 4 0 13 36 0 0 31 0 225 

Trap-nights 480 0 2,640 240 240 0 1,200 720 240 0 240 0 240 240 0 0 480 0 6,960 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


83 


ORDER LAGOMORPHA- Pikas, Hares, and 
Rabbits 

Family Leporidae 

Sylvilagus audubonii arizonae (Allen, 1877) 
Desert Cottontail 

The Desert Cottontail is broadly distributed 
across most of the arid southwestern United States and 
northern Mexico. Like most other rabbits and hares, it 
tends to be crepuscular (Chapman 1999). The biology 
and natural history of this species was summarized by 
Chapman and Willner (1978). In Nevada, the species 
is known only from creosote and sagebrush habitats in 
the southern tip of Nevada in Clark, Nye, Esmeralda, 
White Pine, and Lincoln counties (Hall 1946). Two 
Desert Cottontail (young-of-the-year) were captured 
in PDFAs in Dune (Mesquite) habitat in 2008. 

Additional records. —"At Pahrump Ranch, Indian 
Springs, and St. Thomas, cottontails were even more 
numerous than jack rabbits [sic]” (Burt 1934:423). 

Known museum specimens. —Ash Meadows, 4.8 
mi NW of Devil’s Hole (2, MVZ). 

Lepus californicus deserticola Mearns, 1896 
Black-tailed Jackrabbit 

The Black-tailed Jackrabbit is a conspicuous part 
of the mammalian fauna of the desert Southwest. It is 
known to occur in very diverse habitats from near sea 
level to montane elevations (North and Marsh 1999). 
The biology and natural history of this species was 
summarized by Best (1996). It occurs throughout 
Nevada and is more likely to be seen than other rabbit 
and hare species (Hall 1946). Although Black-tailed 
Jackrabbits were commonly observed on the Refuge, 
none was trapped or collected during field efforts. 

Additional records. —"Jack rabbits [sic] were 
abundant about the farming districts. There were 
hundreds of them at Pahrump Ranch, Indian Springs, 
Corn Creek Station, Las Vegas, and St. Thomas” (Burt 
1934:423). 

Known museum specimens .—None. 


ORDER SORICOMORPHA- Insectivores 
Family Soricidae 

Notiosorex crawfordi crawfordi (Coues, 1877) 
Crawford’s Desert Shrew 

This small-bodied insectivore is considered rare 
in Nevada (Mantooth and Riddle 2005) and has been 
reported from only three southern counties in the state 
(Clark, Nye, and Mineral). A single Crawford’s Desert 
Shrew was captured in a pitfall trap set in Riparian 
Shrubland habitat along the outflow below King’s Pool 
in 2009. This is the first known record of the species 
from the Refuge and only the seventh record of its oc¬ 
currence in Nevada (Manning et al. 2013). The biol¬ 
ogy and natural history of this shrew was reviewed by 
Armstrong and Jones (1972) and Armstrong (1999a). 

Additional records .—None. 

Known museum specimens. —1, TTU. 

ORDER CARNIVORA - Carnivores 
Family Canidae 

Can is latrans mearnsi Merriam, 1897 

Coyote 

The Coyote is a common, widely distributed 
canid of the United States. It is often heard more than 
seen. On the Refuge 8-10 individuals were recorded 
and they were heard nearly every time we were on 
the Refuge after dark. Most sightings were in Alkali 
Shrub-Scrub or Creosote Shrubland. The biology and 
natural history of this predator were reviewed by Bekoff 
(1977, 1999). 

Additional records .—None. 

Known museum specimens .—None. 

Urocyon cinereoargenteus scottii Mearns, 1891 
Common Gray Fox 

The Common Gray Fox is far less common on 
the Refuge than the Coyote. During the 2-year study, 
two sightings were recorded for this canid—one near 


84 


Clyde Jones Memorial Volume 


Crystal Reservoir and another on the north end of the 
Refuge in an Alkali Scrub area. Haroldson (1982) and 
Fritzell (1999) reviewed the biology of this small fox. 

Additional records .—None. 

Known museum specimens .—None. 

Family Felidae 

Puma concolor kaibabensis (Nelson and Goldman, 
1931) 

Mountain Lion 

The Mountain Lion is often associated with 
montane habitat in desert regions. Refuge personnel 
relayed a reported sighting of a Mountain Lion in the 
southeast portion of the Refuge. Several people also 
reported seeing a large cat in the area a few years prior 
to the initiation of this study. Currier (1983) and Beier 
(1999) reviewed the biology and natural history of this 
large felid. 

Additional records .—None. 

Known museum specimens .—None. 

Lynx rufus baileyi Merriam, 1890 
Bobcat 

The bobcat is a fairly common predator in the 
desert Southwest. Refuge staff report several sight¬ 
ings each year. During the study we documented one 
individual crossing the road on the southern end of the 
Refuge. Itwasseeninanareaoflowmesquite. Lariv- 
iere and Walton (1997) and Layne (1999) summarized 
the biology of this nocturnal felid. 

Additional records .—None. 

Known museum specimens .—None. 

Family Mustelidae 

Taxidea taxus berlandieri Baird, 1857 
American Badger 

The American Badger is a semi-fossorial carni¬ 
vore. One of its preferred dietary items is the pocket 


gopher. Long (1973,1999) reviewed the natural history 
and biology of this species. Only one American Badger 
was observed on the Refuge by one of us during the 
2-year study. This animal was seen from a helicopter 
on the north end of the Refuge west of Longstreet 
Spring. In addition, an American Badger was reported 
at the west entrance of the Refuge on Spring Meadows 
Road in March 2010. Finally, two skulls were found 
by Refuge staff, one in sandy habitat on the north end 
of the Refuge, and one in a cattail marsh along Fair¬ 
banks Outflow. 

Additional records .—None. 

Known museum specimens .—None. 

ORDER ARTIODACTYLA- Even-toed Ungulates 
Family Antilocapridae 

Antilocapra americana americana (Ord, 1815) 
Pronghorn 

Pronghorn occur over much of the western and 
southwestern United States, including much of the State 
of Nevada. Although no Pronghorns were observed 
during the field study, there is at least one observation 
of the species from the Refuge: “... we have confirmed 
sightings of pronghorn (1) and mule deer” ... on the 
Refuge (Cristi Baldino, USFWS, Ash Meadows NWR). 
O’Gara (1978, 1999) reviewed the biology of this 
unique artiodactyl. 

Additional records .—None. 

Known museum specimens .—None. 

Family Bovidae 

Ovis canadensis nelsoni Merriam, 1897 
Desert Bighorn Sheep 

The Desert Bighorn Sheep is the state mammal 
of Nevada. Of the three subspecies of desert bighorns 
known to occur in the state, O. c. nelsoni is the smallest 
and palest (Hall 1995). Shackleton( 1985) reviewed the 
biology of this species. Bighorn Sheep were observed 
on the Refuge during this study along the higher and 
lower elevations near Point of Rocks in September 
and October. 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


85 


Additional records .—None. 

Known museum specimens .—None. 

Family Cervidae 

Odocoileus hemionus hemionus (Rafinesque, 1817) 
Desert Mule Deer 

The Desert Mule Deer occurs over most of the 
western United States, including most of Nevada. It is 
a big game animal over much of its range. Although 
no Desert Mule Deer were documented during the 
field study, there is at least one reported observation 
of the species from the Refuge: “...we have confirmed 
sightings of pronghorn and mule deer” ... on the Ref¬ 
uge (Cristi Baldino, USFWS, Ash Meadows NWR, 
personal communication). McCullough (1999) and 
Anderson and Wallmo (1984) reviewed the biology 
of this taxon. 

Additional records .—None. 

Known museum specimens .—None. 

ORDER RODENTIA- Rodents 
Family Cricetidae - New World Mice, Rats, and 
Voles 

Neotoma lepida lepida Thomas, 1893 
Desert Woodrat 

The Desert Woodrat (also known as the Desert 
Packrat) is found in the Great Basin, southern Califor¬ 
nia, and Baja California. It has a vegetarian diet and 
usually constructs rather large nests that may be used 
by many generations of Desert Woodrats (MacMillen 
1999). The biology and natural history of this species 
were summarized by Verts and Carraway (2002). It 
occurs statewide in Nevada, except for the north-central 
part of Nevada. 

A total of 37 Desert Woodrats were taken dur¬ 
ing our trap efforts. These rats tended to be rather 
broadly distributed across the Refuge, and individuals 
were captured in a variety of habitats: Cottontop Dry 


Ridge (n = 10); Emergent Marsh (n = 7); and Alkali 
Shrub-Scrub (n = 7). These three habitats accounted 
for 24 individuals (64.9% of all captures). Only eight 
Desert Woodrats were captured in pitfall traps, seven in 
Riparian Shrubland and one in Dune (Mesquite) habitat. 

Additional records .—None. 

Known museum specimens. —Ash Meadows (5, 
NMNH). 

Onychomys torridus longicaudus Merriam, 1889 
Southern Grasshopper Mouse 

This is a small, territorial, carnivorous rodent 
of the Sonoran and Mojave deserts of North America. 
Insects and scorpions are a main staple of its diet 
(Riddle 1999). The Southern Grasshopper Mouse has 
a shorter white-tipped tail than similarly sized rodents 
found in the same areas. It is most often associated with 
sandy habitats, and its “sand bathing” behavior keeps 
its pelage clean and in prime condition. The biology 
and natural history of this species were summarized by 
McCarty (1975). In Nevada it is known only from the 
southernmost regions. 

Only seven of these carnivorous rodents were 
taken in Sherman live-traps. Four grasshopper mice 
were captured in Dune (Scrub) habitat, and the other 
three individuals were taken in Dune (Mesquite) habi¬ 
tat. Many Southern Grasshopper Mice (n = 25) were 
taken in pitfall traps. Most individuals (n = 11) were 
taken in Dune (Scrub) habitat, whereas seven each 
were captured in Alkali Shrub-Scrub and Dune (Mes¬ 
quite) habitat. All habitats in which grasshopper mice 
were captured had a fine sand substrate as a major 
component. 

Additional records .—None. 

Known museum specimens .—Ash Meadows, 2.5 
mi NW of Devil’s Hole (1, MVZ); Ash Meadows, 4.8 
mi NW of Devil’s Hole (2, MVZ); Ash Meadows (2, 
NMNH). 


86 


Clyde Jones Memorial Volume 


Peromyscus crinitus stephensi Mearns, 1897 
Canyon Deermouse 

The Canyon Deermouse usually is found in habi¬ 
tat that has large amounts of exposed rock, vertical or 
horizontal, with only scant vegetation. Often this is 
the only mammal species living on isolated mesas or 
buttes (Armstrong 1999b). The biology and natural 
history of this species was summarized by Johnson and 
Armstrong (1987). There are three known subspecies 
of the Canyon Deermouse in Nevada (Hall 1946). 

Only five Canyon Deermice were taken in Sher¬ 
man live-traps. All of these mice came from higher el¬ 
evations in rocky areas in Cottontop Dry Ridge habitat. 
No individuals were taken in a pitfall trap. 

Additional records .—None. 

Known museum specimens. —Ash Meadows, 4.8 
mi NW of Devil’s Hole (1, MVZ); Ash Meadows (2, 
NMNH). 

Peromyscus eremicus eremicus (Baird, 1847) 
Cactus Deermouse 

This denizen of desert environs prefers low rocky 
slopes with scattered vegetation and is distributed from 
western Texas to southern California and northern 
Mexico. Its diet consists of plant material and insects. 
The Cactus Deermouse may enter daily torpor or 
seasonal aestivation during the hottest months of the 
summer (Caire 1999). In Nevada, this species is known 
only from the extreme southern tip of the state in Nye, 
Clark, and Lincoln counties (Hall 1946). 

The Cactus Deermouse was third only to Mer- 
riam’s Kangaroo Rat and Western Harvest Mouse in 
terms of individuals taken (n = 267). It was captured in 
nearly every habitat sampled on the Refuge. The four 
habitats with highest numbers of captured individuals 
were Riparian Woodland (n = 48), Emergent Marsh ( n 
= 57), Transitioning Agriculture (n = 18), and Alkali 
Shrub-Scrub (n = 18). The Cactus Deermouse was 
the most frequently taken rodent in pitfall traps (n = 
112, or 49.7% of all captures). Four habitat types had 
especially high numbers of captures: Riparian Wood¬ 
lands, Transitioning Agriculture, Alkali Shrub-Scrub, 


and Ash. Veal and Caire (1979) reviewed the biology 
of this species. 

Additional records. —"Pahrump Ranch” (Burt 
1934:416). 

Known museum specimens .—Ash Meadows, 
4.8 mi NW of Devil’s Hole (2, MVZ); Ash Meadows 
(17, NMNH). 

Peromyscus maniculatus sonoriensis (LeConte, 
1853) 

North American Deermouse 

This deermouse is one of the most ubiquitous 
mice in the United States. It occurs from the Pacific 
Coast to the Atlantic Coast, except for the southeastern 
states, and from southern Alaska and Canada to Mexico. 
It is also one of the more geographically and ecologi¬ 
cally variable species of the genus (Handley 1999). 
In southern Nevada, this deermouse often is restricted 
to riparian habitats (Hall 1946). Two subspecies are 
known to occur statewide in Nevada (Hall 1946). 

No North American Deermice were captured in 
2008. The following year five individuals were cap¬ 
tured. Two were taken in Sherman live-traps, one from 
Transitioning Agriculture and one from Alkali Sink 
habitat. Three individuals were taken in pitfall traps, 
one each from Alkali Shrub-Scrub, Creosote Shrubland, 
and Alkali Meadows habitats. 

Additional records .—None. 

Known museum specimens. —Ash Meadows (10, 
NMNH). 

Reithrodontomys megalotis megalotis (Baird, 1857) 
Western Harvest Mouse 

The Western Harvest Mouse is rather broadly 
distributed over the central grasslands and southwestern 
deserts of North America. It is usually associated with 
thick, mesic vegetation on firm soils (Webster 1999). 
The biology and natural history of this species were 
summarized by Webster and Jones (1982). Reithro- 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


87 


dontomys is somewhat smaller in body size than other 
sympatric peromyscine rodents (Hall 1946). 

Two hundred seventy-five Western Harvest Mice 
were taken in Sherman live-traps. Most individuals (n = 
90) were trapped in Alkali Meadows habitat, frequently 
in areas where the house mouse (Mus musculus) was 
a common inhabitant. Other individuals were taken 
across the Refuge in areas with short dense vegetation, 
especially wet areas (e.g., Wet Meadow, n = 59; and 
Emergent Marsh habitat, n = 44). Western Harvest 
Mice also were taken in pitfall traps (n = 31). Most 
of these individuals (n = 8) were captured in Alkali 
Shrub-Scrub habitat, with the remainder coming from 
nearly all other habitat types, albeit in low numbers. 
This species was not taken in areas devoid of ground 
vegetation (e. g., Creosote Shrubland, Salt Desert 
Scrub, and Cottontop Dry Ridge). 

Additional records. —"Pahrump, Indian Springs, 
Las Vegas, Wheeler Well, Willow and Cold Springs, 
Mormon Well, and St. Thomas” (Burt 1934:416). 

Known museum specimens. —Ash Meadows, 2.5 
mi W of Devil’s Hole (2, MVZ); Ash Meadows (28, 
NMNH). 

Family Geomyidae 

Thomomys bottae centralis Hall, 1930 
Botta’s Pocket Gopher 

Botta’s Pocket Gopher is found throughout the 
southwestern deserts of the United States and northern 
Mexico. This fossorial rodent is highly variable in body 
size and pelage color, and there is marked sexual di¬ 
morphism in many populations across its known range 
(Patton 1999). In Nevada, four species of Thomomys 
are known to occur (Hall 1946). Specimens at NMNH 
were identified as T. b. perpallidus by J. Patton (R. 
Fisher, NMNH, personal communication). From an e- 
mail communication with Dr. Patton, the following was 
received: “Whether or not centralis itself (type locality 
in White Pine Co., eastern Nevada) is a synonym of 
perpallidus awaits analyses of Nevada samples of T. 
bottae .” Further, he stated, “I curated all of the pocket 
gophers in the USNM [United States National Museum] 
collection back in 1993 and at that time believed, and 
still do, that those samples from Nye and Clark coun¬ 


ties that Hall listed as centralis were best considered 
perpallidus .” 

Until the Nevada material has been studied in 
detail, especially using molecular techniques, the 
conservative stance is taken for this report; therefore, 
centralis is retained as the correct subspecific name 
at this time. It is understood that with study, what is 
referred to as centralis in this report may indeed be 
assigned to T. b. perpallidus in the future. 

Only one pocket gopher was trapped during the 
first year of the study. It was taken using a Baker- 
Williams gopher trap in Emergent Marsh habitat adja¬ 
cent to a saltcedar ( Tamarix sp.) removal site next to a 
cattail ( Typha sp.) marsh at Crystal Reservoir. During 
2009 a single young-of-the-year, presumably dispersing 
from natal burrow systems, was taken in a Sherman live 
trap in Alkali Meadows habitat. Four individuals were 
captured in pitfall traps, one each in four habitat types: 
Alkali Shrub-Scrub, Alkali Meadow, Transitioning 
Agriculture, and Emergent Marsh. Evidence of gopher 
burrowing activity commonly was seen on the Refuge. 
Most frequently the burrows and diggings were seen 
in areas with loose, sandy-loamy soils, frequently on 
slightly elevated surfaces such as berms. 

Additional records. —"At Pahrump Ranch, Indian 
Springs, Las Vegas, and St. Thomas” (Burt 1934). 

Known museum specimens. —Ash Meadows, 2.5 
mi W of Devil’s Hole (18, MVZ); Ash Meadows, 4.8 
mi W of Devil’s Hole (11, MVZ); Ash Meadows (41, 
NMNH). 

Family Heteromyidae 

Chaetodipus formosus mojavensis Huey, 1938 
Long-tailed Pocket Mouse 

A saxicolous species, the Long-tailed Pocket 
Mouse occurs in the Great Basin, Mojave, and Colorado 
deserts of western North America. This medium-size 
pocket mouse is often associated with creosotebush, 
shadscale, and various sagebrush species (Geluso 
1999). In Nevada, this mouse is most frequently is 
found in the Lower Sonoran Zone (creosote) and 
lower part of the Upper Sonoran Zone (sagebrush) 
(Hall 1946). 


88 


Clyde Jones Memorial Volume 


A total of 48 Long-tailed Pocket Mice were 
taken in Sherman live-traps during this study. Most 
individuals (n = 24) were taken in Cottontop Dry Ridge 
habitat and Creosote Shrubland (n = 21). Both habitat 
types possess rocky soils or desert pavement. Fourteen 
Long-tailed Pocket Mice were taken in pitfall traps. Ten 
individuals were taken in Creosote Shrubland habitat; 
the other four were taken in Alkali Shrub-Scrub habitat. 

Additional records .—None. 

Known museum specimens .—None. 

Dipodomys deserti deserti Stephens, 1887 
Desert Kangaroo Rat 

The Desert Kangaroo Rat is a large-bodied, 
four-toed, sand dune-dwelling species. It constructs 
an extensive burrow system with numerous entrances 
(Best 1999). The biology and natural history of this 
species was summarized by Best et al. (1989). This 
is the largest and palest of the Dipodomys species in 
Nevada. 

A total of 47 Desert Kangaroo Rats were captured 
during this study, with most individuals taken in areas 
of fine, windblown sand. Twenty-eight individuals 
were captured in Dune (Shrub) habitat, eleven in Dune 
(Mesquite), four in Alkali Sink, three in Alkali Shrub- 
Scrub, and one in Alkali Meadows. None were taken 
in pitfall traps. 

Additional records. —"Pahrump, Indian Springs, 
Com Creek, and St. Thomas” (Burt 1934:415). 

Known museum specimens. —Ash Meadows (39, 
NMNH). 

Dipodomys merriami merriami Mearns, 1890 
MerrianTs Kangaroo Rat 

This four-toed kangaroo rat occupies a rather 
broad range of habitat and soil types. When found with 
congeners, it primarily occurs on rocky soils (Rogers 
1999); however, it also occurs on sandy and clayey 
soils. MerrianTs Kangaroo Rat ranges farther out onto 
alkali flats than other species of Dipodomys (Hall 1946). 


This was the most frequently trapped rodent 
during the study, with a total of 276 individuals cap¬ 
tured. Three habitats accounted for most (n = 210, or 
76%) of the captures: Alkali Shrub-Scrub (n = 84), 
Creosote Shrubland (n = 63), and Dune (Mesquite) ( n 
= 63). Only 12 MerrianTs Kangaroo Rats were taken 
in pitfall traps; nine specimens were captured in Alkali 
Shrub-Scrub habitat and three were captured in Dune 
(Mesquite). 

Additional records .—None. 

Known museum specimens .—Ash Meadows, 
2.5 mi NW of Devil’s Hole (2, MVZ); Ash Meadows, 
4.8 mi NW of Devil’s Hole (1, MVZ). Ash Meadows 
(35, NMNH). 

Dipodomys microps occidentalis Hall and Dale, 
1939 

Chisel-toothed Kangaroo Rat 

The Chisel-toothed Kangaroo Rat is named for 
its flattened chisel-like incisors, which differ from the 
rounded awl-like incisors of other kangaroo rats. It 
has five toes and is somewhat unusual in that it feeds 
primarily on plant leaves rather than on seeds. A key 
plant in its diet is shadscale. The known distribution 
of the Chisel-toothed Kangaroo Rat is the arid Great 
Basin between the Sierra Nevada and Rocky Moun¬ 
tains (Hayssen 1999). The biology and natural history 
of this species were summarized by Hayssen (1991). 
This taxon is found in western and southern Nevada 
(Hall 1946). 

This was the least common of the three kangaroo 
rat species found on the Refuge. Fifteen Chisel-toothed 
Kangaroo Rats were captured during the 2-year study. 
Nine were captured in Alkali Shmb-Scmb, one in Creo¬ 
sote Shrubland, and three in Salt Desert Scrub. Only 
two Chisel-toothed Kangaroo Rats were taken in pitfall 
traps, both in Alkali Shrub-Scrub habitat. 

Additional records .—None. 

Known museum specimens. —Ash Meadows (3, 
NMNH). 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


89 


Perognathus longimembris panamintinus 
Merriam, 1894 
Little Pocket Mouse 

This small heteromyid is fairly common through¬ 
out the southwestern United States. It ranges from 
southern Oregon through most of Nevada, southern 
California, and northern Baja California (French 
1999). In Nevada, this pocket mouse usually is found 
on “...the fine sandy soil in the center of valleys, but 
is found more often, and more abundantly, on firmer 
soils of the slightly sloping margins of the valleys” 
(Hall 1946:358). 

Only seven Little Pocket Mice were captured 
in our Sherman live-traps. Four were taken in Alkali 
Shrub-Scrub and one each was taken in Salt desert 
Scrub, Transitioning Agriculture, and Dune (Mesquite). 
Seven individuals were taken in the pitfall traps, all in 
Alkali Shrub-Scrub habitat. 

Additional records .—None. 

Known museum specimens .—None. 

Family Muridae 
Mus musculus Linnaeus, 1758 
House Mouse 

This invasive, non-native rodent usually is con¬ 
sidered a commensal of humans. It generally is found in 
or near man-made structures, although some survive in 
native habitats. The House Mouse frequently is found 
in the same habitat (mesic areas with dense vegetation, 
where it is considered abundant) as harvest mice (Hall 
1946). These moist habitats are a central component of 
the Refuge’s conservation efforts for the Ash Meadows 
Montane Vole. Competition with native mice may 
contribute to the possible demise of the Montane Vole. 

One hundred thirty of these mice were taken 
in Sherman live traps during the 2-year study. Two 
habitats were particularly noted to harbor House 
Mice; Wet Meadow (n = 43) and Alkali Meadow ( n 
= 37). This species usually was taken in areas with 
rather dense ground cover on moist soils. Only three 
House Mice were captured in pitfall traps, one each, in 


Alkali Shrub-Scrub, Creosote Shrubland, and Riparian 
Shrubland habitats. 

Additional records. —"Pahrump Ranch” (Burt 
1934:422). 

Known museum specimens. —Ash Meadows, 4.8 
mi NW of Devil’s Hole (2, MVZ). 

Family Sciuridae 

Ammospermophilus leucurus leucurus (Merriam, 
1889) 

White-tailed Antelope Squirrel 

The White-tailed Antelope Squirrel is a heat- 
tolerant diurnal sciurid. Unlike other squirrels, this 
desert rodent is active throughout the year. In addition 
to eating plant material, it also consumes insects and 
small rodents (Hafner 1999). The biology and natural 
history of this species was summarized by Belk and 
Smith (1991). This squirrel can be found in areas 
with sandy soils, as well as on firmer, even rocky, soils 
(Hall 1946). 

Fifteen White-tailed Antelope Squirrels were 
taken in this 2-year study. Most individuals (n = 12) 
were captured in Alkali Shrub-Scrub habitat (one in 
a pitfall trap), whereas three were taken in Creosote 
Shrubland; individuals occasionally were seen along 
roadways on the Refuge while driving through various 
habitat types. 

Additional records .—None. 

Known museum specimens. —Ash Meadows (16, 
NMNH). 

Xerospermophilus tereticaudus tereticaudus Baird, 
1857 

Round-tailed Ground Squirrel 

Round-tailed Ground Squirrels are found pri¬ 
marily in sandy areas of relatively flat desert habitat, 
especially near mesquite, creosotebush, and saltbush. 
The biology and natural history of this desert rodent 
was reviewed by Ernest and Mares (1987). Like the 


90 


Clyde Jones Memorial Volume 


White-tailed Antelope Ground Squirrel, this species 
is heat tolerant and may remain active when tempera¬ 
tures exceed 45°C (113°F) (Ernest 1999). In Nevada, 
the Round-tailed Ground Squirrel is known only from 
southern Clark and Nye counties (Hall 1946). 

There are four specimens from the Death Valley 
Biological Survey of 1891 from Ash Meadows. This 
species was captured only once during our 2-year study 
on the Refuge, in Alkali Sink habitat. 

Additional records. —None. 

Known museum specimens .—Ash Meadows (4, 
NMNH). 

Mammals of Possible Occurrence 

ORDER CARNIVORA 
Family Canidae 

Vulpes macrotis arsipus Elliot, 1903 
Kit Fox 

This small fox has been united systematically and 
taxonomically with the Swift Fox, which is known to 
occur farther to the east and north. Most authorities 
now consider the Kit Fox to be a separate species. Mc- 
Grew (1979) and Thacker and Flinders (1999) reviewed 
the biology and natural history of this taxon. No Kit 
Foxes were observed on the Refuge, but a single kit 
fox was seen crossing the road between the Refuge and 
Pahrump, Nevada. 

Family Mephitidae 

Mephitis mephitis estor Merriam, 1890 
Striped Skunk 

We had no encounters with the Striped Skunk 
while conducting field work at the Refuge. Wade-Smith 
and Verts (1982) and Bixler (1999) reviewed the biol¬ 
ogy and natural history of this skunk. 

Spilogale gracilis gracilis Merriam, 1890 
Western Spotted Skunk 

The Western Spotted Skunk frequently is as¬ 
sociated with rocky, somewhat vertical habitat. This 


species was not encountered during the field work on 
the Refuge. Crooks (1999) and Verts and Carraway 
(2001) reviewed the biology and natural history of 
this small skunk. 


Family Procyonidae 

Bassariscus astutus nevadensis Miller, 1913 
Ringtail 

This procyonid frequently is associated with 
rocky outcrops and cliffs, often near permanent water 
sources. It was not observed during the study at Ash 
Meadows NWR. Poglayen-Neuwald and Toweill 
(1988) and Baker (1999) summarized the biology of 
this species. 

ORDER RODENTIA 
Family Cricetidae 

Microtus montanus nevadensis Bailey, 1898 
Montane Vole 

The Montane Vole is a common and widely dis¬ 
tributed rodent of the intermountain western United 
States. It is found in a variety of mesic habitats, 
including grasslands, wet meadows, and streamside 
vegetation (Jannett 1999). Anderson (1959) revised 
and summarized the biogeography and taxonomy of 
North American Montane Voles. Hoffmann and Koeppl 
(1985) discussed the zoogeography of New World mi¬ 
crotines. The biology and natural history of this species 
was summarized by Sera and Early (2003). In Nevada, 
the species occurs mostly in montane habitats in the 
northern half of the state. Montane Voles are active 
throughout the day during the growing seasons, but 
peak activity is at dusk and dawn (Jannett 1999; Sera 
and Early 2003). The vole frequently is associated with 
habitats dominated by graminoid vegetation and can 
be found in well-drained, arid, and wet habitats (Getz 
1985). Montane Voles are semi-fossorial and their 
nests can be found both aboveground and belowground; 
however, brood nests always occur belowground. In 
addition to the nests they dig, male Montane Voles 
may also use and occupy gopher burrows. Currently, 
15 subspecies of Montane Moles have been described 
(Anderson 1959; Hall 1981; Sera and Early 2003). 
Most subspecies of the Montane Vole have not been 
studied thoroughly due to their limited distribution 


Manning and Heaney—Mammals of Ash Meadows National Wildlife Refuge 


91 


(Sera and Early 2003). Hall (1946) and Mantooth 
and Riddle (2005) both reported six subspecies of the 
Montane Vole in Nevada, one of which was the Ash 
Meadows Montane Vole. 

Bailey (1898) first described the Ash Meadows 
Vole as Microtus nevadensis. Bailey (1900) later 
revised the genus {Microtus). Hall (1935) suggested 
that it was a subspecies of the montanus- group and 
applied the trinomial Microtus montanus nevadensis. 
This taxon (known from 30 specimens and minimal 
field data) was collected at “... a big salt marsh below 
Watkins Ranch, Ash Meadows, Nye County, Nevada” 
(Hall 1946:549). 

The Nevada Natural Heritage Program lists 
the Ash Meadows Montane Vole as “SH” state rank, 
which denotes that the species is known from histori¬ 
cal records and could be rediscovered (Mantooth and 
Riddle 2005). Historically, the Ash Meadows Montane 
Vole was known to occur on the Refuge. We were not 


able to locate this vole during our field trapping ef¬ 
forts. In addition to our live-trap study (this report), 
we sampled select mesic habitats (on the northern part 
of the Refuge). None was found. The species appears 
to be extirpated from the Refuge. 

Additional records. —None. 

Known museum specimens. —Ash Meadows, 
4.8 mi NW of Devil’s Hole (13, MVZ); Ash Meadows 
(17, NMNH). 

Family Muridae 

Rattus norvegicus norvegicus (Berkenhout, 1769) 
Norway or Brown Rat 

This introduced rodent is found statewide in 
Nevada. The Norway Rat is a commensal with man 
and is generally found in and around human habitation. 
No individuals were taken at the Refuge. 


Discussion 


The Refuge supports a diverse terrestrial mam¬ 
malian fauna dominated by a few species. In this study, 
the Cactus Deermouse, the Western Harvest Mouse, and 
Merriam’s Kangaroo Rat dominate the rodent commu¬ 
nity in terms of relative abundance. The mammalian 
community of the Refuge appears to be representative 
of the region, at least in terms of species diversity. Of 
the 18 native, non-volant small mammal species (sorici- 
morphs, lagomorphs, and rodents) whose distribution 
includes the Refuge, 17 (94%) were documented during 
this study. Only the Ash Meadows Montane Vole was 
not documented to occur there during our study. Other 
species not observed on the Refuge were carnivores, 
and one of these (Kit Fox) was observed just outside 
the Refuge boundary. The Ringtail, Western Spotted 
Skunk, and Striped Skunk were not observed during 
the study. Moreover, the methods employed during 
this study targeted primarily smaller ground foraging 
mammals. 

The success of future trapping efforts for small 
mammals using Sherman live-traps could be maxi¬ 
mized by seasonal timing. Trapping efforts conducted 
in early autumn would coincide with peak rodent 


populations, following spring and summer recruitment. 
Additional efforts could also be implemented to target 
the Desert Shrew (e.g., by installing much smaller 
pitfall traps placed at higher densities). Such efforts 
could provide better insight to the range and density 
of the Desert Shrew. 

Based upon the results of this study, it appears 
likely that Ash Meadows Montane Vole has been extir¬ 
pated from the Refuge. This conclusion is supported 
by the lack of success by other researchers since the 
vole last was reported in 1933. Moreover, because most 
potential habitat in the Ash Meadows marsh complex is 
located on the Refuge, it is unlikely that populations ex¬ 
ist in areas not searched during this or previous studies. 

Competition with introduced rodent species also 
might have played a role in inhibiting Ash Meadows 
Montane Vole populations. We found that the House 
Mouse was abundant in areas of potential vole habitat 
on the Refuge. Some authors have suggested that 
invasion by introduced species has negative popu¬ 
lation-level consequences for native rodent species 
(Simberloff 2009). 


92 


Clyde Jones Memorial Volume 


We were anxious to document the mammals that 
eke out a living in this region. The Mojave Desert is in 
one of the hottest and driest, habitats in North America. 


What we found helps us define or understand the niche 
of these desert-adapted species. 


Acknowledgments 


We acknowledge the following members of the 
Ash Meadows National Wildlife Refuge staff for their 
assistance at the Refuge: Sharon McKelvey (Refuge 
Manager) and Cristi Baldino (Wildlife Biologist). We 
thank Ben Chen, Jeff Enright, Jeremy Hull, T. Wayne 
Schwertner, and Tara Raabe. Special thanks to the 
“vegetation crew” of BIO-WEST, Inc., for all the help 
with botanical aspects of field study. All trapping 


efforts were conducted under BIO-WEST’S Nevada 
Department of Wildlife’s scientific collection permit 
(permit no. S-29822) and the USFWS’s Special Use 
Permit. Frank D. Yancey, II and Larry L. Choate read 
and improved an earlier version of this paper. The 
authors are responsible for making editing suggestions 
and any mistakes. 


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Anderson, S. 1959. Distribution, variation, and relationships 
of the montane vole, Microtus montanus. University 
of Kansas Publications, Museum of Natural History 
9:415-511. 

Anderson, A. E., and O. C. Wallmo. 1984. Odocoileus hemionus. 
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San Marcos, Texas 78666 

rwmanning@ grandecom.net 


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Martin R. Heaney 

Bio-West Inc. 

1018 Frost Drive 
Rosenberg, Texas 77471 
mheaney @ bio-west, com 


A Resurvey of Bats at Dinosaur National Monument 

Michael A. Bogan and Tony R. Mollhagen 


Abstract 

Between 1982 and 1990, biologists associated with the U.S. Fish and Wildlife 
Service (FWS) conducted inventories for mammals, including bats, at Dinosaur National 
Monument (DINO) in Colorado and Utah. These studies by FWS were the first sys¬ 
tematic attempts to survey for bats throughout the monument. In 2008, park managers 
asked for a resurvey of the resident bat fauna. The basic premise was to re-visit localities 
where bats had been captured between 1982 and 1990 and again use mist nets to assess 
the general distribution and abundance of bats on the monument and determine if there 
were any obvious trends in species occurrences, numbers, or distribution. In the original 
study, we netted a total of 60 nights and captured 468 individuals of 14 species at 26 
localities. Our average capture rate was 7.8 individuals of three species per sampling 
event. In 2008-2009, we netted a total of 51 nights and captured 909 individuals of 
15 species at 23 localities. The average capture rate was 17.8 bats of 4.4 species per 
netting event. Examination of the two data sets, separated by more than two decades, 
reveals far more similarities than differences. Recent work yielded almost twice as 
many bats as the former study, yet the inventory of species is essentially unchanged. 
In both time periods, a large majority of the total number of bats was taken at higher 
elevations, and male bats of two-thirds of the species dominated the composition at 
higher elevations. From a faunal perspective, the 15 resident bat species at DINO can 
be divided into three classes: abundant (> 10% of total captures); common (2-10% 
of captures); and uncommon (< 2% of captures). Bats that were abundant at DINO 
included Myotis evotis, M. volans, M. yumanensis, Lasionycteris noctivagans, and 
Eptesicus fuscus. Bats that were common included M. californicus, M. ciliolabrum, 
M. thysanodes, Lasiurus cinereus, Corynorhinus townsendii, and Antrozous pallidus. 
Uncommon species at DINO included M. lucifugus, Parastrellus hesperus, Euderma 
maculatum, and Tadarida brasiliensis. There is one record for Nyctinomops macrotis 
from the monument, but we did not capture one in either time period. Most species 
were in the same classes in the earlier study, suggesting the bat fauna is relatively stable 
over time. We consider most of the resident species to be stable in numbers given the 
results of the two studies. For two species, M. ciliolabrum and M. evotis, the capture 
numbers indicate they might be increasing slightly. For three others, M. lucifugus, L. 
cinereus, and E. fuscus, the numbers suggest that there might be some evidence of a 
downward trend. 

Key words: bats, Colorado, Dinosaur National Monument, distribution and 
abundance, population, resurvey, trends, Utah 


Introduction 

Originally established in 1915 to protect paleon- Monument (DINO) was expanded in 1938 to include 
tological resources on 32 hectares, Dinosaur National the Green and Yampa river corridors. The present 


97 


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85,446-hectare monument is rich in geological and 
biological resources, and its singular landscapes of 
river canyons and upland benches are unique among 
lands managed by the National Park Service (NPS). 
The monument includes among its biological resources 
a possible 16 species of bats. Bats are subjects of 
conservation concern due to habitat destruction, use of 
insecticides, low reproduction rates, and vulnerability 
due to colonial roosting habits (O’Shea and Bogan 
2004). Western bats in particular also may be affected 
by land management practices such as grazing, logging, 
lowering of water tables, and destruction of reservoirs. 
In addition, ongoing changes in the area surrounding 
DINO may conceivably impact bats or resources upon 
which they depend. Such changes include: perceived 
decline of cottonwood gallery forests; grazing, oil 
and gas development; increases in recreational use; 
invasion of exotic species; and attempts to remove 
one such exotic (tamarisk, Tamarix). Finally, ongoing 
global changes, such as climate warming, may affect 
bats (Humphries et al. 2002; LaVal 2004; Rebelo et 
al. 2010). 

Between 1982 and 1990, biologists associated 
with the Fish and Wildlife Service (FWS) conducted 
inventories for mammals, including bats, in the 
monument. Initially, we surveyed the river corridors 
in DINO in support of a reserved water rights case that 
later proceeded through the court system. Subsequent 
survey efforts were expanded to upland areas in the 
monument, many of which proved rich in numbers of 
bats. Although some of the oldest records of bats in 
Colorado are from the area around DINO (e.g., Cary 
1911, summarized by Armstrong 1972), these studies 
by FWS were the first systematic attempts to survey 
for bats throughout the monument (Bogan et al. 1983, 
1988). The work resulted in the capture of more than 
450 individuals from 26 localities on or very near the 
park; many captured individuals were saved as voucher 
specimens and deposited in the Biological Survey 
Collections, Fort Collins, CO [now located in the 
Museum of Southwestern Biology (MSB), University 
of New Mexico], Efforts by other investigators 
also resulted in a variety of new information for the 
monument and Moffat County, including work by 
Freeman (1984), who surveyed throughout the state 
of Colorado but worked extensively in Moffat County, 
Navo et al. (1992), who conducted the first acoustic 
survey at the park, and Storz (1995), who obtained 
interesting new data on Euderma maculatum (Spotted 


Bat). Although most information comes from the 
Colorado (Moffat County) portion of the monument, 
there also is information from the Utah (Uintah County) 
side. In 2008-2009, we conducted a “resurvey” effort 
of bats at DINO. The basic premise was to re-visit 
localities where bats had been captured between 
1982 and 1990, assess the general distribution and 
abundance of bats, and determine if there were changes 
in species occurrences, numbers, or distributions on 
the monument. 

The state of Colorado is known to have 19 species 
of bats (Chiroptera) (Armstrong et al. 1994; Fitzger¬ 
ald et al. 1994; Hayes et al. 2009). Of these 19, two 
are species that have recently invaded from the east, 
Lasiurus borealis (Eastern Red Bat) and Perimyotis 
subflavus (Tricolored Bat); these two species are not 
known to occur on the western slope of Colorado. 
Another species, Idionycteris phyllotis (Allen’s Big- 
eared Bat) was recently confirmed from acoustic data in 
southwestern Colorado (Hayes et al. 2009). For much 
of western Colorado there are records for 16 species. 
Utah is known to have 18 species, 16 of which occur 
in northeastern Utah (Uintah Co.). Only L. blossevil- 
lii (Western Red Bat) and I. phyllotis are absent from 
Uintah County. Overall, there are verified records of 
16 species from DINO, including an enigmatic record 
of Nyctinomops macrotis (Big Free-tailed Bat) from the 
parking lot at the Dinosaur Quarry on the Utah side of 
the monument (Table 1). 

We suspect that it is the complex geological, 
topographical, and biological features of the park that, 
in part, produce the diverse bat fauna of DINO (e.g., 
Humphrey 1975). Additionally, the characteristics of 
each species, coupled with their biogeographic history 
and location on the Colorado Plateau, contribute to 
this rich fauna. Indeed, the bat faunas of DINO and 
Canyonlands National Park (CANY), about 190 km 
S W DINO, for example, are remarkably similar (Bogan 
et al. 2006). Each park has 16 species but whereas 
there are records for Myotis lucifugus (Little Brown 
Myotis) but not for I. phyllotis at DINO, the converse 
is true for CANY. Another well-known area, the Henry 
Mountains in south-central Utah, is known to have 16 
species as well (Mollhagen and Bogan 1997). 

At the time of Durrant’s (1963) surveys for mam¬ 
mals in DINO, shooting was the common method of 
obtaining specimens of bats in the field. As no shooting 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


99 


Table 1. Bats (Chiroptera) known to occur in the region of Dinosaur National Monument 
(Durrant 1952; Hall 1981; Fitzgerald et al. 1994; Mollhagen and Bogan 1997; Bogan et al. 
2006; O’Shea et al. 2011). Scientific and common names follow Baker et al. (2003) except 
for Parastrellus, which follows Hoofer et al. (2006). The acronyms for the species names 
are those used in tables elsewhere in this report. 


Scientific Name 

Acronym 

Common Name 

Myotis californicus 

Myca 

California Myotis 

M. ciliolabrum 

Myci 

Western Small-footed Myotis 

M. evotis 

Myev 

Long-eared Myotis 

M. lucifugus 

Mylu 

Little Brown Myotis 

M. thysanodes 

Myth 

Fringed Myotis 

M. volans 

Myvo 

Long-legged Myotis 

M. yumanensis 

Myyu 

Yuma Myotis 

Lasiurus cinereus 

Laci 

Hoary Bat 

Lasionycteris noctivagans 

Lano 

Silver-haired Bat 

Parastrellus hesperus 

Pahe 

Canyon Bat 

Eptesicus fuscus 

Epfu 

Big Brown Bat 

Euderma maculatum 

Euma 

Spotted Bat 

Corynorhinus townsendii 

Coto 

Townsend’s Big-eared Bat 

Antrozous pallidus 

Anpa 

Pallid Bat 

Tadarida brasiliensis 

Tabr 

Brazilian Free-tailed Bat 

Nyctinomops macrotis 

Nyma 

Big Free-tailed Bat 


was allowed in the monument, Durrant and his crew 
obtained no bats. However, Durrant listed 14 species 
of bats that he presumed occurred on the monument, 
based on his knowledge of mammals of the Colorado 
Plateau (e.g., Durrant 1952). Indeed, all 14 of these 
species are known today, along with M. californicus 
(California Myotis) and the Big Free-tailed Bat. Arm¬ 
strong (1972), in his study of Colorado mammals, found 
specimens in collections for just eight species of bats 
in Moffat County: M. yumanensis (Yuma Myotis), M. 
evotis (Long-eared Myotis); M. volans (Long-legged 
Myotis); M. ciliolabrum (Small-footed Myotis); Ep- 
tesicus fuscus (Big Brown Bat); L. cinereus (Hoary 
Bat); Corynorhinus townsendii (Townsend’s Big-eared 
Bat); and Antrozous pallidus (Pallid Bat). Armstrong 
et al. (1994) had records of 14 species from Moffat 
County, excluding the molossids Tadarida brasiliensis 
(Brazilian Free-tailed Bat) and N. macrotis. Distribu¬ 


tion maps in Fitzgerald et al. (1994) likewise indicated 
the presence of 14 species in Moffat County, again 
excluding only the molossids. There are records for 
both molossids from DINO, although only from the 
Utah side of the monument. 

Four of the 16 species of bats occurring within 
DINO are state species of concern for Colorado and/ 
or Utah. These are C. townsendii in both states, and 
M. thysanodes (Fringed Myotis), E. maculatum, and N. 
macrotis in Utah. In Colorado, all bats are protected 
as nongame animals. Management Policies (2006:45) 
state that the NPS will inventory native species that are 
of special management concern (such as rare, declin¬ 
ing, sensitive, or unique species and their habitats) and 
will manage them to maintain their natural distribution 
and abundance. 


100 


Clyde Jones Memorial Volume 


The primary objective of this resurvey was to 
attempt to document the occurrence of at least 90% of 
the bats documented in the previous studies at DINO 
by means of a two-year field effort using mist nets, as 
in the original field efforts. New, pertinent records 


and specimens were examined as necessary. The work 
involved re-visiting sites netted in 1982-1990, setting 
nets, collecting data on bats, and spending about 30 
days per year in the field over two years. 


Methods 


Most methods used in the study were consistent 
with original methods and with methods currently 
approved by bat biologists for such work (e.g., Kunz 
and Kurta 1988; Kunz et al. 2009) and are detailed 
in a written capture and handling protocol approved 
annually by the Institutional Animal Care and Use 
Committee (IACUC) at the University of New Mexico. 
All bats captured were released unharmed as rapidly as 
possible. No voucher specimens were taken nor were 
there any bat mortalities during our work in 2008-2009. 
We obtained a research permit from the monument 
(DINO-2008-SCI-0013) to allow the survey. Although 
annual reports were provided to DINO during our work 
from 1982 to 1990, no overall synthesis of the data was 
ever completed. To provide a database for comparison 
with the recent work, we entered the earlier data into a 
spreadsheet. We included data from specimens in the 
MSB collected by us during this time period as well 
as capture numbers from field notes that we deemed 
reliable. As a part of this phase, we also re-examined 
the museum specimens to verily identifications. 

Mist-net surveys .—Mist nets (Avinet Inc., 
Dryden, NY) were deployed across and around bod¬ 
ies of water (e.g., Haystack Rock Reservoir) and in 
perceived flyways (e.g., Hog Canyon, Split Mountain) 
usually, but not always, at sites and in patterns similar to 
the original surveys. Lengths of mist nets ranged from 
3 to 20 m and numbers of nets deployed on any single 
evening varied from one to five, depending on the area 
and shape of the body of water. Mist nets were set up 
shortly before sunset and tended for several hours until 
activity declined. Nets were never left untended. Ef¬ 


fort was recorded as total horizontal meters of standard 
nets deployed and net nights (total nets x nights). We 
attempted to approximate the number of nights in the 
original sampling; we tallied 60 separate netting events 
in 1982-1990 and 51 in 2008-2009. 

We removed bats from nets immediately fol¬ 
lowing capture and recorded time of capture, species, 
sex, reproductive condition, and any miscellaneous 
comments on standardized field data sheets and then 
released the bats. We later summarized the data from 
field data sheets in Excel spreadsheets for tabulation 
of data, calculation of the number of species and 
individuals captured, relative abundance of species 
(percent of all bats captured), and prevalence value 
(the percent occurrence among all possible dates and 
locations sampled). For the purpose of comparison, 
the same treatment was given to data from the earlier 
survey. Copies of field data sheets were provided to 
the monument. 

Location data .—Each of the 23 sites where we 
sampled in 2008-2009 was given a waypoint name 
and a more descriptive name. Geographic coordinates, 
elevation, and estimated position error were acquired 
for each locality with Garmin GPS units set to record 
coordinates in decimal degrees using NAD27 as a 
datum. The NAD 1927 datum was used because that 
is the datum for USGS quad maps we employed for 
orientation. Elevations obtained by GPS units were 
reconciled against the USGS quad maps and when 
there was a discrepancy between sources, values were 
interpolated. 


Results and Discussion 

During the two-summer study, we set out 2,480 (Table 2). In 2008 we made three trips to DINO, 
meters of bat nets and accrued a total 195 net nights encompassing 83 total person-days of travel, to recon- 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


101 


Table 2. Level of effort at all sites in 2008-2009 showing number and size of nets at each netting event, total 
horizontal meters of net deployed at each event, and net nights (nets x nights). 


Location 

Date 

Chew Reservoir 

8-Aug-08 

Snow Reservoir 

9-Jul-09 

Buffham Reservoir 

8- Jul-08 

9- Jul-08 

Massey Pond 

l-Jul-08 

Massey Reservoir 

30-Jun-08 

18-Jul-09 

Bear Draw Reservoir 

5-Aug-08 

17-Jul-09 

Dry Woman Reservoir 

6-Jul-08 

4-Aug-08 

17-Jul-09 

Haystack Rock Reservoir 

4- Jul-08 

5- Jul-08 

3- Aug-08 

4- Jun-09 

16-Jul-09 

Ely Creek 

14-Jul-09 

Vermillion Creek 

7-Jul-08 

Hog Canyon 

l-Jun-08 

Pool Creek Ranch 

12-Aug-08 

Morris Ranch 

31-May-08 

2- Jul-08 

1-Aug-08 

10-Aug-0 8 

3- Jun-09 

17-Jun-09 

Big Joe Campground 

9- Jun-09 

10- Jun-09 

Pool Creek Petroglyphs 

3-Jun-08 


3m 


2 

1 


6m 


1 


1 


2 

3 

1 

1 


2 

3 


10m 14m 20m 

3 
5 

1 3 

1 1 2 

1 1 

2 2 

1 3 

4 

1 1 1 

2 1 
4 

2 1 2 

1 1 2 

1 3 

4 

5 

1 4 
1 

2 1 
3 

1 2 

3 2 


4 

4 

5 

2 2 

1 

1 


Total 

Meters 

60 

too 

74 

64 

40 

60 

74 

80 

50 

48 

80 

74 

64 

74 

80 

too 

94 

26 

18 

48 

66 

56 

82 

80 

80 

too 

68 

26 

30 

12 


Net 

Nights 

3 
5 

4 
4 

3 

4 
4 
4 
4 

3 

4 

5 
4 
4 

4 

5 
5 
3 
3 

3 

4 

4 

5 
4 

4 

5 

4 

5 
5 
4 


3 


1 


102 


Clyde Jones Memorial Volume 


Table 2. (cont.) 


Location 

Date 

3m 

6m 

10m 

14m 

20m 

Total 

Meters 

Net 

Nights 


10-Jul-08 

3 


6 

3 


11-Aug-08 

2 


1 

1 

38 

4 


15-Jun-09 

2 

1 

1 

1 


34 

5 


8-Jul-09 

2 

1 


10 

3 

Pot Creek 

12-Jul-09 


1 

1 

2 

64 

4 

Harding Hole 

ll-Jun-09 

2 

2 

1 


26 

5 


12-Jun-09 

4 


1 


18 

5 

Pool Creek at Echo Park 

2-Jun-08 


4 

1 


34 

5 


ll-Jul-08 

1 

2 


14 

3 


6-Aug-08 


1 


6 

1 


16-Jun-09 

1 

2 

1 


24 

4 

Cub Creek 

3-Jul-08 


3 


18 

3 


2-Aug-08 


3 


18 

3 


7-Jun-09 


4 


24 

4 


10-Jul-09 


3 


18 

3 

Laddie Park 

13-Jun-09 


1 

2 

2 

78 

5 

Rippling Brook 

13-Jul-09 

1 

2 


1 


28 

4 

Jones Hole Campground 

14-Jul-09 


2 


12 

2 

Split Mountain 

9-Aug-08 


1 


14 

1 


6-Jun-09 


2 

1 

48 

3 


10-Jul-09 


2 


1 

40 

3 


Total 

2,480 

195 


noiter netting sites and to net bats. We were in DINO 
during 29 May to 5 June, 30 June to 12 July, and 1-13 
August for a total of 68 person-days. During this time 
we netted a total of 27 nights at 15 different locali¬ 
ties (Table 2) for a total of 95 net-nights. Inclement 
weather precluded netting on several occasions. Four 
of the 15 localities were new ones that were not netted 
in the earlier work. We netted the following localities 
in 2008: Chew Reservoir (1 time); Buffham Reservoir 
(2 times); Massey Pond (1, new); Massey Reservoir 
(1); Bear Draw Reservoir (1, new); Dry Woman Res¬ 
ervoir (2); Haystack Rock Reservoir (3); Vermillion 


Creek (1); Hog Canyon (1, new); Morris Ranch (4); 
Pool Creek Ranch (1); Pool Creek Petroglyphs (3); 
Pool Creek at Echo Park (3); Cub Creek (2); and Split 
Mountain (1, new). 

In 2008, we captured 517 bats of 15 species 
(Table 3), more individual bats than we captured in the 
original survey. Earlier, we averaged 7.8 bats of 3.0 
species per night, whereas in 2008 we averaged 19.1 
bats of 4.8 species per night. Included in the 2008 
averages are two nights when we set nets but captured 
no bats (Hog Canyon and Split Mountain). The best 


Table 3. Localities, dates, and bat species captured in Dinosaur National Monument in the summer of2008. Data are arranged by highest elevation and earliest 
netting date. The columns on the right side indicate the total number of bats and total bat species captured by locality and date. Summaries at the bottom include 
the total individuals of each species (Sum), the percentage that species comprises of the 517 total bats captured (% Total Bats), and the percentage of times at 
least one individual of that species was captured among the 27 separate combinations of sampling locations and dates (% Prevalence). The identities of the 
acronyms for the 15 bat species are given in Table 1. Descriptions and coordinates of the capture localities are in locality accounts available from the authors. 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


103 


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104 


Clyde Jones Memorial Volume 


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one-night captures came from Haystack Rock Reser¬ 
voir (96 and 56), Dry Woman Reservoir (51), Morris 
Ranch (47), Pool Creek at Echo Park (39), Haystack 
Rock Reservoir (36), and Buffham Reservoir (33). Our 
earlier efforts convinced us that Haystack Rock Reser¬ 
voir was an important resource for bats and based on 
our netting there in July 2008 this continued to be true. 
Even in August 2008, when the pond was mostly mud 
and the surrounding area showing the effects of large 
numbers of cattle, nets set around the periphery still 
captured bats attracted by an insect hatch from the mud. 

We captured all species of bats known from 
DINO except for N. macrotis , known only by a salvaged 
specimen from the Quarry area but presumed to be a 
member of DINO’s bat fauna. Among the other three 
species of concern, numbers from the original surveys 
versus those from 2008 are: M. thysanodes 5, 15; E. 
maculatum 5, 5; and C. townsendii 14, 19. Great cau¬ 
tion should be used in extrapolating from these num¬ 
bers, but they seem to demonstrate that these species 
of concern are still a part of the bat fauna of DINO and 
in roughly the same or greater numbers as in the past 
based on our netting data from 2008 (Table 3). 

The most abundant species we captured in 2008 
were: E. fuscus, 84; M. evotis , 74; M. yumanensis, 
68; L. noctivagans, 68; A. pallidus, 67; and M. volans, 
46. Total captures of the other species ranged from 
one (T. brasiliensis ) to 33 (M ciliolabrum). Also of 
note was the capture of four Canyon Bats (Parastrellus 
Hesperus) in 2008. This species is on the edge of its 
range at DINO and previously was known by only two 
individuals from the monument. 

We made two trips to DINO in 2009, with an em¬ 
phasis on netting at some of the sites in the river corri¬ 
dors that were netted in 1982 (Table 4). Our total travel 
was 70 person-days and we worked on the monument 
3-18 June and 8-19 July for a total of 56 person-days. 
During our time on the monument, we netted 24 nights 
at 17 different sites for a total of 100 net-nights. Spe¬ 
cifically, we netted at Snow Reservoir (1 night, new), 
Massey Reservoir (1), Bear Draw Reservoir (1), Dry 
Woman Reservoir (1), Haystack Rock Reservoir (2), 
Split Mountain (2), Morris Ranch (2), Cub Creek (2), 
Pool Creek at Echo Park (1), Pool Creek Petroglyphs 
(2), Big Joe Campground (2, Yampa River), Harding 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


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Elev. Total Total 

(m) Location Date Myca Myci Myev Mylu Myth Myvo Myyu Laci Lano Pahe Epfu Euma Coto Anpa Tabr Bats Species 


106 


Clyde Jones Memorial Volume 


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Hole (2, Yampa River), Laddie Park (1, Yampa River), 
Pot Creek (1, Green River), Rippling Brook (1, Green 
River), Jones Hole Campground (1, Green River), and 
Ely Creek (1, at Jones Hole, Green River). 

In 2009, we netted a total of 392 individuals of 
15 species (Table 4). On average, we captured 16.3 
individuals of four species each night we netted, down 
slightly from the 2008 averages (19.1; 4.8). Included 
in these averages are three nights when we captured 
no bats. The most productive localities were Haystack 
Rock Reservoir (101 individuals), Dry Woman Reser¬ 
voir (71), Bear Draw Reservoir (65), Haystack Rock 
Reservoir (38), and Snow Reservoir (27), all upland 
sites south of the Yampa River Corridor. Among the 
river sites, Laddie Park (7) and Harding Hole (6) were 
the most productive. In general, our captures at the 
river sites were disappointingly low, perhaps because 
of some combination of low temperatures in June, net¬ 
ting sites cluttered by vegetation, and abundant water. 
Among the species of concern, we captured seven M. 
thysanodes, six E. maculatum, and 10 C. townsendii, 
numbers similar to captures in 2008. Again, these 
species were captured at roughly the same proportion 
as in years past. 

The most frequently captured species in 2009 
included 119 L. noctivagans, 64 E.fuscus, 47 M. evotis, 
38 M. volans , and 31 M. ciliolabrum. Total captures of 
other species ranged from one (T. brasiliensis) to 23 (M. 
yumanensis, Table 4). The most notable difference be¬ 
tween captures in the two years was the near-doubling 
of numbers of L. noctivagans, the large reduction in 
numbers of A. pallidus, fewer M. evotis and M. yuma¬ 
nensis, and the slight reduction in numbers of E.fuscus 
and M. volans. Although there are multiple biases in 
using mist nets, we are more inclined to suspect that 
many, or all, of the between-year differences are the 
result of more mundane factors including somewhat 
lower temperatures in June 2009 as well as random 
chance in our choice of when to net a given site. For 
example, on 2 June 2008 we captured a large number of 
female M. yumanensis, likely from a maternity colony, 
at Pool Creek at Echo Park. Similarly, we captured 28 
A. pallidus at the Morris Ranch one night (Table 3). 
Although we netted both sites in 2009, it was at dif¬ 
ferent times and we did not encounter such numbers. 
Adult male L. noctivagans are common to abundant 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


107 


in midsummer at DINO and at times may account for 
up to 50% of all bats captured on the Yampa Bench 
(Tables 3,4). The large number caught in 2009 reflects 
this abundance. 

We and our associates netted on or very near the 
park in six different years: June and July 1982; August 
1985; July 1987; August 1988; June 1989; and July 
1990. In 1982, all localities were in either the Yampa or 
Green river canyons, areas where we suspected that bat 
netting was not very productive due to the abundance of 
available water. Most netting from 1985 to 1990 was 
conducted primarily at upland sites, including several 
productive sites on the Yampa Bench and in or near 
northern areas of the monument. The exact times of our 
visits were generally determined by a variety of factors 
(e.g., work scheduled elsewhere) rather than by phe¬ 
nomena such as moon phase or expected temperatures. 

Between 1982 and 1990, we captured 468 in¬ 
dividuals of 14 species (Table 5); no molossids were 
taken. Our average capture rate was 7.8 individuals of 
3.2 species per sampling event. On eight of 60 (13%) 
separate combinations of locations and dates, we cap¬ 
tured no bats. We captured relatively high numbers of 
individuals at Massey Reservoir (21.4 average), Big 
Joe Campground (19.5), Dry Woman Reservoir (18), 
Haystack Rock Reservoir (15.4), Canyon Overlook 
(11.3), Pool Creek Petroglyphs (11), and Cub Creek 
(6.8). Most of the productive sites ranged in elevation 
from 1,974 m to 2,346 m, although Big Joe, Pool Creek 
Petroglyphs, and Cub Creek ranged only from 1,566 
m to 1,600 m. 

In terms of average number of species, the most 
productive netting sites were Massey Reservoir (7 
species), Haystack Rock Reservoir (6.7), Canyon 
Overlook (4.7), Dry Woman Reservoir (4.5), Big Joe 
Campground (4.5), Pool Creek Petroglyphs (4), and 
Cub Creek (2.8, Table 5). At most other sites we usually 
captured 1-3 species per night. Typically, the best of 
these sites were above 1,974 m, had moderately large, 
still waters that allowed many species to use them, and 
pools were often isolated from other water resources. 
The small pools upstream from Big Joe Campground 
on the Yampa River were an exception, perhaps due 
to an abundant insect resource or nearby favorable 
roosting sites. 


Among the species of concern, we captured seven 
M. thysanodes (1.5% of total bats), five E. maculatum 
(1.1%), and 14 C. townsendii (3%) between 1982 and 
1990 (Table 5). By comparison, the most abundant 
species were E. fuscus (112, 24%), L. noctivagans 
(69, 15%), M. volans (55, 12%), M. yumanensis (52, 
11%), and M. evotis (41,9%), andL. cinereus (41,9%). 
Most of these more abundant species are known to be 
common at higher elevations, although M. yumanensis 
also are common at lower elevations (e.g., Pool Creek 
at Echo Park). It is worth noting that some species 
were uncommon or missing among our captures. For 
example, between 1982 and 1990 we took only two P. 
hesperus (0.4%) and no molossids (Table 5) although 
we obtained a salvaged T. brasiliensis taken in 1985 
from the Green River Housing Area on the Utah side 
of the monument. The N. macrotis was salvaged from 
the Dinosaur Quarry (Utah, Uintah Co., R23E, T4S, 
Sec. 26) by W. Dye on 9 December 1996 and is now 
in the MSB. 

Among the 26 sites where we worked from 1982 
to 1990 (Table 5), we were able to net at 17 of these sites 
in 2008-2009 (Table 6). The nine sites we did not net 
included two where bats were shot (Massey Camp and 
Haystack Rock on Yampa River); among the seven net- 
table sites were three that were dry (Bufifham Place, Old 
Bassett Cabin, Massey Troughs), two sites on private 
property (Five Springs, Canyon Overlook), one on the 
river could not be scheduled (Alcove Brook), and one 
was deemed too large and deep to net (Cottonwood 
Creek near Canyon Overlook). Of the seven nettable 
sites, six were upper elevation sites (above 2,134 m) 
but accounted for less than 10% of all captures from 
1982 to 1990 (n = 45). Only one of these sites, Canyon 
Overlook, was very productive; we netted a total of 34 
bats of nine species there in July 1990. 

Among the historic sites, the 17 we did net in 
2008-2009 were Chew Reservoir, Buffham Reservoir, 
Massey Reservoir, Bear Draw Reservoir, Dry Woman 
Reservoir, Haystack Rock Reservoir, Vermillion Creek, 
Pool Creek Ranch, Ely Creek, Morris Ranch, Big Joe 
Campground, Pool Creek Petroglyphs, Pot Creek, 
Harding Hole, Pool Creek at Echo Park, Cub Creek, 
and Rippling Brook. We added six new sites to com¬ 
pensate for those no longer available or usable. These 
included Snow Reservoir, Massey Pond (dry in 2009), 
Hog Canyon, Laddie Park, Jones Hole Campground, 


Table 5. Localities, dates, and bat species captured in Dinosaur National Monument, 1982-1990. Data are arranged by highest elevation and earliest netting 
date. The columns on the right side indicate the total number of bats and total bat species captured by locality and date. Summaries at the bottom include 
the total individuals of each species (Sum), the percentage that species comprises of the 468 total bats captured (% Total Bats), and the percentage of times at 
least one individual of that species was captured among the 60 separate combinations of sampling locations and dates (% Prevalence). The identities of the 
acronyms for the 15 bat species are given in Table 1. Descriptions and coordinates of the capture localities are in locality accounts available from the authors. 


108 


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Clyde Jones Memorial Volume 


and Split Mountain (Table 6). Technically, Bear Draw 
Reservoir was a new site. It was less than a mile from 
the original site in Bear Draw yet it provided more 
surface area for drinking by bats than the original pool. 

Overall, we captured almost twice as many bats 
(909) in the recent work (Table 6) as in the original 
surveys (468; Table 5). We captured 17.8 bats of 4.4 
species per netting event in the recent work compared 
to 7.8 bats of 3.0 species per event in the original 
survey. Among higher-elevation sites, the most pro¬ 
ductive sites were: Haystack Rock Reservoir (65.4 
bats/night); Dry Woman Reservoir (45.3); Bear Draw 
(39.5); Snow Reservoir (27 on one night); and Bulfham 
Reservoir (22). The more productive lower-elevation 
sites were: Morris Ranch (14.8); Pool Creek at Echo 
Park (14.3); Pool Creek Petroglyphs (8.4); and Cub 
Creek (8.0). Many of these sites were productive in 
the earlier study as well (Dry Woman, Haystack, Pool 
Creek Petroglyphs, and Cub Creek), although Massey 
Reservoir and Big Joe Campground yielded far fewer 
captures. Several performed better than in previous 
visits (Bulfham Reservoir, Bear Draw, Morris Ranch, 
and Pool Creek at Echo Park). 

The reasons for increased capture rates are un¬ 
known, as we did not attempt to increase our numbers 
and in fact netted fewer nights (60 vs. 51). One dif¬ 
ference between the two periods is that in the earlier 
work we also conducted rodent surveys and prepared 
specimens, so perhaps our efforts on bats were diluted 
to some extent. Countering this is that in much of 
the earlier work we left nets set throughout the night, 
a practice we no longer conduct or condone. In the 
recent work, we also made some attempt to schedule 
our work at the time of the dark (new) moon, because 
in our experience bat captures increase under these 
conditions. However, in 2009 we were less able to do 
this due to river trip schedules. There is the possibility 
that we deployed more long (20 m) nets in 2008-2009 
but we cannot reconstruct data from the original study 
to facilitate this comparison. Finally, we suspect we 
might be very slightly better at netting bats as we have 
had 20-plus years of additional experience. 

During our work in 2008-2009 (Table 6), we 
netted a total of 15 species at DINO. We believe this 
number represents the number of resident species at 


the monument and consider N. macrotis as occasion¬ 
ally present. For most of the 15 species, we obtained 
evidence of reproduction (data available from authors). 
In terms of numbers of species per site, the higher- 
elevation sites demonstrating greater diversity were: 
Dry Woman Reservoir (10 species/night); Haystack 
Rock Reservoir (8.6); Buffham Reservoir (8.5); and 
Snow Reservoir (7, one night). Lower-elevation sites 
tended to have fewer species: Morris Ranch (4.6); 
Pool Creek Petroglyphs (4.4); and Cub Creek and 
Pool Creek at Echo Park (3.5 per site). Again, several 
of these were diverse in 1982-1990 (Dry Woman, 
Haystack, Pool Creek at Petroglyphs, and Cub Creek), 
although Massey Reservoir and Big Joe failed to meet 
the diversity observed in the earlier survey. 

It seems axiomatic that sites that have high diver¬ 
sity (and numbers) are important to the bats that occur 
there and are an important source of food, water, and 
perhaps social interactions. We recommend that these 
sites be placed on a list of “high-priority” sites for wild¬ 
life at DINO and consideration be given to protecting 
them. This does not mean that sites with low diversity 
are not important as well; they may have lower diversity 
because not all species can obtain a given resource there 
(e.g., too small for large bats to maneuver over but an 
excellent site for smaller bats). We do not know why 
two sites (Massey Reservoir and Big Joe Campground) 
that were previously productive in numbers or species 
were not as productive in 2008-2009. In the case of 
Massey, it was our subjective opinion that there were 
many more cattle present than in 1990. Perhaps there 
have been detrimental changes in insect populations, 
roosting sites, or water quality, or maybe it was merely 
a consequence of timing. Additional work at the site 
might clarify this situation. In the case of Big Joe 
Campground, there is the possibility that our memo¬ 
ries and notes failed us and we did not net far enough 
upstream in Starvation Valley in 2009. 

In terms of numbers of the species of concern, 
we were reassured by our captures of three species. 
We caught over three times as many M. thysanodes 
and twice as many E. maculatum and C. townsendii as 
in the previous study. Among these three species, the 
percent of total captures went up for M. thysanodes 
but stayed about the same for the other two species. 
Prevalence values for all three went up, suggesting they 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


115 


were more widespread than before. We did not net any 
N. macrotis so their status at DINO remains unclear. 

One clear similarity between the two periods was 
the preponderance of bats captured at high-elevation 
sites. Between 1982 and 1990, we captured 311 (66%) 
bats at sites above 1,800 m and 157 (34%) at sites 
below that elevation. There were 13 sites both above 
and below 1,800 m. In 2008-2009, eight sites were 
above 1,829 m and 15 sites were below that elevation 
(Table 6). Nonetheless, sites above 1,800 m yielded 
657 (72% of the total) bats in 2008-2009, whereas 252 
(28%) came from the 15 lower sites. Are bats really 
more common at higher elevations in the monument or 
are these numbers a result of other factors? 

For example, did we spend more time or effort 
at the higher elevations? In terms of net-nights, in 
2008-2009 we expended 69 (35.3%) net-nights above 
1,800 m and 126 (64.6%) net-nights below that eleva¬ 
tion. In terms of effort, we deployed 1,112 m (48.5%) 
of net at sites above 1,829 m elevation and 1,178 m 
(51.4%) of nets below that elevation. Thus, we had a 
slightly greater effort at lower elevations than at higher 
sites, a fact influenced to some degree by revisiting 
the sites on the rivers. We could not reliably calculate 
net-nights or net feet deployed for the earlier work but 
assume it was similar. Thus, time or effort would not 
seem to account for greater numbers of captures at the 
high-elevation sites. 

Are there inherent differences between the 
low- and high-elevation sites? Here, there may be 
some definite trends. DINO is a land of large, rolling, 
sagebrush-covered benches at the upper elevations 
dissected by multiple small drainages and canyons that 
ultimately, for the most part, drain into the canyons 
of the Yampa and Green rivers. Cattle grazing is an 
historic land-use pattern in the upper areas; although 
some grazing occurs at lower elevations (e.g., Cub 
Creek), it seems less common and intense. One of 
the consequences of grazing the benches has been 
the construction and maintenance of “stock ponds” or 
reservoirs for livestock. Although such reservoirs are 
sometimes controversial, and we know of instances 
where land-management agencies have destroyed such 
impoundments, many bat biologists see them as a vital 
resource for bats, especially in areas where the water 


table has dropped (e.g., Mollhagen and Bogan 1997; 
Rabe and Rosenstock 2005; Jackrel and Matlack 2010; 
Chambers et al. 2011). 

These scattered, moderate to large, reservoirs on 
the benches are desirable sites for bat netting as bats 
seem to congregate at such places. It is possible that 
such nutrient-rich stock ponds increase aquatic insect 
biomass, and thus increase forage for bats. They tend to 
be larger in size than most lowland sites and thus have 
more surface area for foraging bats. They also may be 
closer to roosting sites. Virtually all lower-elevation 
sites are smaller in size. Many are located on small 
streams that are often subject to flash flooding (which 
may affect in situ insect biomass) or are sites where we 
could not place nets over water (e.g., Morris Ranch and 
Laddie Park) due to interference by vegetation. In turn, 
most netting sites along the rivers are compromised 
by the abundance of water where bats can forage or 
drink, a phenomenon that may dilute concentrations 
of bats. Whatever the biological reasons may be for 
apparent concentrations of bats at the high-elevation 
sites, they are clearly sites that biologists can use to 
obtain information on bats. 

Are there detectable trends among bats taken at 
the higher elevation sites? During our earlier work, 
we had the impression that we captured more males 
than females at high-elevation sites. Our database for 
2008-2009 gave us an opportunity to examine if this 
might be true. We again used an elevation of 1,800 m 
in elevation to separate low and high elevation sites. 
Although this division is somewhat arbitrary, there is 
a clear separation among our study sites with 15 sites 
ranging from 1,459 m (Split Mountain) to 1,639 m (Ely 
Creek), each differing from the adjacent site by 30-60 
m in elevation. The eight high-elevation sites begin 335 
m above the Ely Creek site at 1,974 m (Haystack Rock 
Reservoir) and range up to 2,263 m (Chew Reservoir), 
each differing from adjacent sites by 30-90 m elevation. 

Our captures of bats at high-elevation sites indeed 
reveal a preponderance of males over females (Table 
7). We netted 511 males (80%) and only 127 females 
(20%). Conversely, at lower sites the numbers were 
more similar although we netted slightly more females 
(151, 59%) than males (103,41%; Table 8). Numbers 
in these tables are slightly different than those shown 


116 


Clyde Jones Memorial Volume 


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netting date. For each net event the number of males captured is given first, then the number of females. The combined total number of bats shown 
captured in tables 7 and 8 is slightly less than the total number shown in Table 6. A few bats escaped before the sex could be determined. The 
identities of the acronyms for the 15 bat species are given in Table 1. 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


117 


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Table 8. (cont.) 


118 


Clyde Jones Memorial Volume 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


119 


in Tables 3-6, because Tables 7-8 exclude bats that we 
identified upon capture but that escaped before all data 
could be gathered. 

Myotis evotis and M. volans , known to be com¬ 
mon at higher elevations (Fitzgerald et al. 1994), had 
the largest numbers of females taken at the higher sites 
(28 for both species; Table 7), presumably reflecting 
this habitat bias. We were surprised by the relative 
abundance of female M. yumanensis (19) at the higher 
sites. This is a species that typically is more common 
near open water (Barbour and Davis 1969) and per¬ 
haps the large reservoirs on the benches attract them 
to these sites. Another anomaly was the number of 
reproductive female C. townsendii that we captured 
at Snow Reservoir on the cool night of 9 July. When 
these bats began to drink and forage over the pool we 
were initially surprised, but we soon concluded they 
had been attracted by a large accumulation of small, 
white moths over the reservoir itself. We observed 
several individuals capture moths. The predilection of 
L. noctivagans to be common at higher-elevation sites 
was mentioned previously. 

Only two species exceeded 28 female captures at 
low-elevation sites (Table 8). At the lower sites we took 
45 female M. yumanensis, more in keeping with known 
habitat predilections of this species, and 29 female E. 
fuscus, a species we often regard as an “elevational” 
generalist. However, in 2008-2009, female E. fuscus 
were more abundant at lower elevations. We also took 
23 female A. pallidus, mostly at the Morris Ranch in 
early August (Table 8). 

At low-elevation sites, where males comprised 
41% of the captures, the numbers of captured males 
exceeded the numbers of captured females in six of 15 
species (Table 9). Conversely, at high-elevation sites, 
where males made up 80% of the catch, captured males 
exceeded captured females in 10 of 15 species (Table 
9). At the higher sites, females outnumbered males 
only for M. lucijugus, M. yumanensis, C. townsendii, 
and T. brasiliensis (n = 1). One species, P. hesperus, 
had equal captures. 

There are compelling reasons for male and female 
bats to occupy different areas, roosts, or habitats during 
the summer (Cryan et al. 2000). Cooler temperatures 


at higher elevations may cause bats to enter torpor. In 
much of the montane West, pregnant females seem to 
select low-elevation habitats as it is important that they 
maintain a constant body temperature to allow either 
embryos or dependent young to develop in the narrow 
window of time before food supplies begin to decline. 
As a part of this strategy, females tend to form maternity 
colonies in sites where their body heat warms the roost. 
By contrast, males are common at higher elevations 
where their strategy is to roost solitarily and enter torpor 
during the day. Among the bats at DINO, females in 
general are more common at lower elevations whereas 
males clearly are more common at higher elevations. 
Female M. evotis and M. volans are an exception to this 
generalization and we assume they are either making 
nightly migrations from low- to high-elevation sites or 
they are roosting at the upper elevations. Both species 
seem adapted to higher elevations. Telemetry studies 
would shed light on the roosting habits of female bats 
at the monument. 

About 37% (98/267) of the females we captured 
in 2008-2009 showed some form of reproductive 
activity: pregnancy, lactation, or sign of recent lacta¬ 
tion (post-lactation). Our netting schedule each year 
precluded any attempt to monitor reproductive cycles in 
detail. The presumption is that most females are preg¬ 
nant early in the summer, but it can be difficult during 
this time to discern definite signs of pregnancy. The 
earliest date we noted gravid bats was 4 July in 2008 
and 16 July in 2009. We captured lactating females 
from 11 July 2008 and 14 July 2009 and as late as 5 
August in 2008. We netted 32 flying young-of-the- 
year but only in 2008, beginning on 3 August and as 
late as 12 August that year. We probably did not work 
late enough in summer 2009 to catch young. During 
the two years, we tallied 113 reproductive males (vas 
deferens swollen and intruding into the uropatagium); 
the earliest dates were 4 July in 2008 and 16 July in 
2009. We continued to net reproductive males until we 
left the monument each year (10 August in 2008 and ca. 
17 July in 2009). Species-specific data on reproduction 
are available from the authors. 

Our data suggest that low temperatures in June 
2009 likely affected bat captures. In June 2008, the 
average temperature when we started a netting event 
was 23° C and ending temperature was 15° C; in June 


120 


Clyde Jones Memorial Volume 


Table 9. Numbers of males and females for low- and high-elevation sites for all bats captured 2008-2009. High 
sites (n = 8) are those above 1,800 meters are shown on the left side of the table. Low sites (n = 15) are below 
that elevation. Data presented are the numbers of each sex of each species and the percent each sex represents 
of the total captures of each species. Bats that escaped before sex determination are not shown. The identities 
of the acronyms for the 15 bat species are given in Table 1. 


Species 


Low Sites 


High Sites 


Males 

% Male 

Females 

% Female 

Males 

% Male 

Females 

% Female 

Myca 

4 

26.7 

11 

73.3 

7 

87.5 

1 

12.5 

Myci 

4 

66.7 

2 

33.3 

49 

92.5 

4 

7.5 

Myev 

12 

60.0 

8 

40.0 

72 

72.0 

28 

28.0 

Mylu 

0 


0 


2 

20.0 

8 

80.0 

Myth 

4 

26.7 

11 

73.3 

9 

81.8 

2 

18.2 

Myvo 

8 

36.4 

14 

63.6 

35 

55.6 

28 

44.4 

Myyu 

13 

22.4 

45 

77.6 

13 

40.6 

19 

59.4 

Laci 

4 

80.0 

1 

20.0 

24 

96.0 

1 

4.0 

Lano 

11 

100.0 

0 

0.0 

172 

100.0 

0 

0.0 

Pahe 

0 


4 

100.0 

1 

50.0 

1 

50.0 

Epfu 

12 

29.3 

29 

70.7 

98 

97.0 

3 

3.0 

Euma 

0 


0 


6 

60.0 

4 

40.0 

Coto 

1 

25.0 

3 

75.0 

3 

12.0 

22 

88.0 

Anpa 

29 

55.8 

23 

44.2 

20 

80.0 

5 

20.0 

Tabr 

1 

100.0 

0 


0 


1 


Total 

103 


151 


511 


127 


2009, starting temperatures were about 4° C lower (19° 
C); ending temperatures were approximately the same 
as in 2008. In June 2008, we captured an average of 
almost 15 bats/night over four nights. In June 2009, 
we averaged less than half that number (7 bats/night), 
even with more nights of effort and a good catch at 
Haystack Rock Reservoir. Three of the 2009 locali¬ 
ties were river sites, accounting for five of 12 nights 
of effort and this may have influenced our captures. It 
is not clear how the presence of a moon in early June 
2009 affected these captures. 

Results during the other three capture periods 
were roughly similar with starting temperatures above 
23° C and ending temperatures around 15-20° C. 
Nonetheless, July 2009, the period with the highest 


average start and end temperatures, had the highest 
average capture rate (25.5 bats/night). Both July efforts 
included productive nights at Haystack Rock Reservoir 
(96 and 101 bats), although the 2009 effort also had 
good nights at Bear Draw and Dry Woman reservoirs. 
In our only August effort (2008) our best capture num¬ 
bers were in the dark of the moon. 

To look for additional patterns we ranked all 
sites by number of bats captured and then looked at 
various attributes of the sites. Not surprisingly all the 
Yampa Bench (upland) sites rank in the top 20 sites 
in number of captures. Additionally, the top eight 
sites, all with captures greater than 39/night, were all 
made when no moon was present. During nights that 
we characterized as “no moon” we captured a total of 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


121 


661 (73%) bats, whereas on nights with some or full 
moon we captured 248 bats. Interestingly, three of 
five nights when we caught no bats also had no moon, 
although all five nights were in river or riparian, not 
upland, sites. Sites we characterized as “upland,” but 
excluding the Morris Ranch, accounted for 657 (72%) 


of all bats captured. Sites characterized as “riparian,” 
“river,” and the Morris Ranch accounted for just 252 
bats. Other variables, such as cloud cover and presence 
of large trees, also probably influence capture rates but 
perhaps in a more subtle fashion. 


Conclusions 


In terms of species diversity, the two time periods 
are very similar. From 1982 to 1990, we captured 14 
species in nets and obtained a fifteenth, a salvaged T. 
brasiliensis. In 2008-2009 we captured all 15 species 
in nets. One of the least common species in the earlier 
work was P. hesperus. We have two specimens from 
that period (one of which was shot), but in 2008-2009, 
we netted a total of seven P. hesperus. In the recent 
study we also netted two T. brasiliensis (including one 
in Colorado). Both of these species are on or near the 
margin of their respective geographic ranges. These 
slight increases suggest to us that the two species are 
clearly surviving in the area, and may be slowly becom¬ 
ing more abundant (Bogan and Cryan 2000; Genoways 
et al. 2000). Other species on or near the edge of their 
ranges include M. californicus, known only to the 
north from adjacent Sweetwater County, Wyoming; M. 
yumanensis , known by us from Big Horn, Lincoln, and 
Sheridan counties, Wyoming (also see Buskirk 2016); 
and E. maculatum and A. pallidus, whose ranges are 
attenuated to the east although more expansive to the 
south, west, and north (Reid 2006). 

When we began this work we had some expecta¬ 
tions that we would capture a N. macrotis as there is a 
salvaged specimen from the Quarry. We are familiar 
with this species from our work in New Mexico and 
Utah and have found reproductive females to be mod¬ 
erately common in some places. We have observed 
their roosts in slickrock canyon walls with large, 
incised cracks and have netted them over long, linear 
pools in streambeds in isolated canyons. The species 
may occur at DINO but it is possible that we did not 
net in locations where they occur. Alternatively, they 
may be absent because some resource is not present 
or in short supply. Bogan and Cryan (2000) report an 
isolated individual (MSB 122221) from West Gros 
Ventre Butte near Jackson, Wyoming, but the northern¬ 


most maternity roosts with females and young are at 
the latitude of Moab, Utah (Bogan, unpubl.). For the 
present, we are inclined to conclude that N. macrotis 
occurs occasionally at DINO. 

A comparison of data from both studies (Table 
10) suggests that in general, bat species and numbers 
are doing well at DINO. Although we caught about 
twice as many bats in the recent study, we do not be¬ 
lieve that bats are twice as abundant as they were in 
1982-1990. For most species we did capture more 
individuals, but there are exceptions. For example, we 
took slightly fewer M. californicus and M. lucifugus 
and about three-fourths the number of L. cinereus. 
Given the relatively small sample sizes, these differ¬ 
ences may not be real. Alternatively, possible reasons 
for the declines are that we may have misidentified a 
few M. californicus as M. ciliolabrum', we were less 
successful at Massey Reservoir where M. lucifugus was 
common in the earlier study; and perhaps we simply 
were not in the right spot at the right time to net more 

L. cinereus. For all three species, the proportion that 
each species represented out of total bats also declined 
by about 50%. Interestingly, their general occurrence or 
distribution on the monument, as measured by percent 
total captures and percent prevalence, remained about 
the same (Table 10). We consider the three species to 
be somewhat less common but as widespread as before. 

Among those species for which we captured 
more individuals, a few increased by factors of three 
to six times. We captured almost three times as many 

M. evotis and L. noctivagans, four times as many M. 
ciliolabrum , and six times as many A. pallidus. It is 
difficult to ignore the possibility that these numbers do 
not represent some real increase in numbers of these 
species (Table 10). Geluso and Geluso (2012) demon¬ 
strated, for example, that bat abundance can fluctuate 


Table 10. A comparison of all bats captured in the periods 1982-1990 and 2008-09. Data tabulated for each period, for each species, 
are the total captures, the percent of total captures, and the percent prevalence of occurrence. The rightmost columns are a comparison 
of the populations as expressed as by the percent of total captures and the percent prevalence for each species. The last column is a 


122 


Clyde Jones Memorial Volume 


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CN 

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187 

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148 


29 

78 

CN 


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1982-90 


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Total 

!h 

G 

G 

24 

IN 

of- 

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IN 

55 

52 

Gt 

69 

CN 

CN 

in 

Gt 

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ro 

o 


u 


t/3 


_QD 

"o 

<D 

G 

CZ2 

Myca 

Myci 

Myev 

Mylu 

Myth 

Myvo 

Myyu 

Laci 

Lano 

Pahe 

Epfu 

Euma 

Coto 

Anpa 

Tabr 


Bogan and Mollhagen—Bats of Dinosaur National Monument 


123 


greatly with precipitation. Nonetheless, such an ap¬ 
parent increase might have resulted from a few very 
successful nights of netting, such as for L. noctivagans 
at Haystack Rock and A. pallidus at Morris Ranch. 
The same may be true for M. evotis, as we had two or 
three very successful nights for that species. However, 
for M. ciliolabrum, the general pattern is different as 
we had many nights of fair to moderate captures (3-8 
animals). The proportion each species made of the total 
catch also increased, as did the prevalence value for all. 
These four species may represent the best evidence that 
some species have increased in numbers. There is the 
possibility that such increases result from subtle habitat 
changes not noticed by us. 

Another suite of species increased by about one 
and one-half to two times in abundance (Table 10). 
These species include M. volans, M. yumanensis, 
E. fuscus, E. maculatum, and C. townsendii. Total 
numbers of each of these species in 2008-2009 vary 
from 11 to 91 and some caution is advised in dealing 
with smaller samples (e.g., E. maculatum ). But, for 
some of these species (e.g., C. townsendii, E. fuscus, 
and M. yumanensis ) we again had a night or two of 
exceptional captures that influenced total numbers. 
Interestingly, the percent each species represented of 
the total remained about the same or decreased slightly. 
Prevalence values increased in most cases, markedly 
so in one case (M. yumanensis ), and decreased in one 
case ( E. fuscus; Table 10). We are persuaded that the 
small change in proportions and the slight increase in 
prevalence suggest these species are healthy at DINO. 

From a faunal perspective, we subjectively di¬ 
vided resident bat species at DINO into three classes: 
abundant (> 10% of total captures), common (2-10% 
of captures), and uncommon (< 2% of captures). Bats 
that are abundant at DINO include M. evotis, M. volans, 
M. yumanensis, L. noctivagans, and E. fuscus. Bats that 
are common include M. califomicus, M. ciliolabrum, 
M. thysanodes, L. cinereus, C. townsendii, and A. pal¬ 
lidus. Uncommon species at DINO are L. lucifugus, 
P. hesperus, E. maculatum, and T. brasiliensis. Most 
species were in the same categories for the earlier work, 
suggesting the fauna is relatively stable over time. The 


exceptions are slight differences in percentages for M. 
evotis, M. thysanodes, and L. lucifugus. 

In many ways, we found DINO unchanged after 
all these years and, reassuringly, the bat fauna of the 
monument still seemed healthy. Nonetheless, we urge 
that our current results be interpreted cautiously at this 
juncture. Why is DINO so “good” for bats? Our guess 
would be that it is a combination of the presence of good 
foraging and roosting sites, coupled with a fortuitous 
distribution of water sources, especially in the uplands. 
The relative paucity of visitors (and their activities) to 
large, wild parts of the monument also may play a role, 
as may the general lack of intensive agriculture near 
the monument. Grazing continues to be a visible land 
use in and near the monument and it was our subjective 
opinion that in some areas, in spite of what appeared to 
be a year of good winter precipitation in 2007-2008, 
the effects of grazing pressure were more obvious than 
during our earlier work. 

Bats at DINO face a variety of local pressures, 
but like all organisms they also face global threats. In 
particular, most climate scientists are convinced that 
global warming is occurring. How would warming 
temperatures affect bats at DINO? Depending on the 
degree of warming, the summer habits of many bats, 
especially males that use torpor at upper elevations, 
could change. How warmer temperatures would affect 
female bats in the summer at DINO is more conjectural 
but presumably some of them would move upward in 
elevation. We know almost nothing about the winter¬ 
ing habits of most bats at DINO other than that some 
(L. noctivagans, L. cinereus, T. brasiliensis) migrate 
to other, warmer, areas where food remains available, 
whereas the other species probably make, at most, 
local migrations to hibernacula to overwinter. Again, 
depending on the degree of warming, some hibernacula 
might become too warm to be used by bats. Although 
this study provides two benchmarks across time com¬ 
paring bat assemblages in one geographic region, the 
data will be informative in comparing future bat assem¬ 
blages, especially in light of threats such as white-nose 
syndrome, should it spread west across the Great Plains. 


124 


Clyde Jones Memorial Volume 


Acknowledgments 


Our work at DINO would not have been possible 
without the interest and cheerful assistance of Cindy 
Heyd, Resource Management Specialist, DINO. Cindy 
assisted with paperwork, ran interference for us, con¬ 
tacted landowners, became familiar with the bats of 
Dinosaur, removed bats from nets, and helped record 
data. Dave Worthington from Capitol Reef National 
Park, an old friend and knowledgeable bat biologist, 
was of great help during our second trip in 2008 and 
Libby Nance, who understands the importance of in¬ 
ventories and monitoring on national parks, assisted us 
on our third trip in 2008. In 2009, Cindy Heyd again 
was instrumental in helping us accomplish our work. 
Cindy arranged our two boat trips, helped with myriad 
details (especially menus), and sacrificed a knee for us 
at Warm Springs Rapid. Cindy Ramotnik assisted us 
on both river trips in 2009, volunteering 27 person- 


days, and we appreciate her willingness to work hard 
and her ability to stay dry. Dave Worthington returned 
in 2009 and accompanied us on the Green River trip 
and, as always, we are grateful for his assistance, good 
humor, and culinary contributions. Pete Williams and 
John Whinery were our boatmen on the Yampa River 
trip in June, and Turner DuPont, Jessie Pierson, and 
Ethan Johnson were the boatpersons on the Green River 
trip in July. We clearly could not have visited the sites 
on the river without their able assistance. Anne Elder 
assisted us with the DINO Research Permit in early 
2008. Finally, we acknowledge Steve Petersburg who 
first introduced us to the beauty of the monument and 
the rewards of working there. Funding for the original 
surveys in 1982 was provided by Dick Weisbrod (NPS) 
and Clyde Jones (FWS). 


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PO Box 2452 
Corrales, NM 87048 
mbogan @ unm. edu 


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Tony R. Mollhagen 

4524 64th S. 

Lubbock, TX 79414 
shellbadger@ sbcglobal net 


Distribution Records and Reported Sightings of the White-nosed Coati 
(Nasua narica) in Texas, with Comments on the Species’ Population and 

Conservation Status 

David J. Schmidly, John Karges, and Robert Dean 


Abstract 

The status of the White-nosed Coati (Nasua narica) in Texas has been an enigma 
for decades. Herein, we review documentation of the historical records as well as numer¬ 
ous reports of recent sightings, very few of which are “confirmed” with specimens or 
photographs; most come from amateur naturalists and must be considered “anectodal.” 
Long-term mammal surveys by professional mammalogists and camera trap studies 
from within the range of the coati in Texas have not produced any specimens or other 
documentation of the species. Almost all of the sightings are of solitary individuals 
from within 100 miles of the borderlands and/or of released pets. The most numerous 
sightings are from Big Bend National Park just across the river from the Maderas del 
Carmen/Sierra del Carmen Mountains of northern Coahuila, Mexico, where a small 
breeding and permanently established population has now become established. We know 
of no evidence that a breeding population of wild coatis now live in Texas, although a 
recent conservation corridor established along the borderlands could eventually result 
in the species expanding its range into the State. 

Key words: conservation, distribution, Nasua narica , Texas, White-nosed Coati 


Introduction 


White-nosed Coatis (Nasua narica ) are procyonid 
carnivores closely related to raccoons and ringtails. 
They are often indiscriminately called coatimundis, 
though the latter term properly refers only to solitary 
adult males. The White-nosed Coati (hereafter simply 
coati) ranges from Panama through Mexico. It reaches 
the limits of its breeding range in the United States 
(U.S.), where it has been recorded in southeastern 
Arizona, southwestern New Mexico, and southwestern 
Texas. The species is uncommon or at low abundance 
throughout its U.S. distribution and especially Texas, 
where it is thought to be rare. The breeding range of 
N. narica in the U.S. extends from the A nim as Moun¬ 
tains in southwestern New Mexico to the Baboquivari 
Mountains in south-central Arizona, and north to the 
Gila River (Kaufmann et al. 1976). 


Although coatis superficially resemble rac¬ 
coons, they have a more slender, flexible snout and a 
long, slender tail that is often carried erect (Kaufmann 
1987). They are intermediate in size between raccoons 
and ringtails, with adults typically weighing 4-6 kg 
(9-13 pounds) and having a total length of 85-134 cm 
(33.5-53 inches), about half ofwhich is tail. Generally, 
female coatis are slightly smaller than males. The color, 
usually chocolate brown, varies from pale sandy brown 
to reddish to almost black. Indistinct dark tail-rings, a 
white snout, broken white-eye rings, and a yellowish 
wash on the chest and shoulders are among the most 
distinguishable characteristics. 

Coatis are thought to be the largest native animal 
to invade the U.S. Borderlands in the last 135 years 


127 


128 


Clyde Jones Memorial Volume 


(Gehlbach 1981). Their northward movement ini¬ 
tially corresponded with post-pluvial times in a fashion 
similar to that of the nine-banded armadillo (Dasypus 
novemcinctus). There is a lack of pluvial fossil records 
for both of these mammals, and they do not appear north 
of the Border until historic times or at archeological 
sites no earlier than a few thousand years Before Present 
(B.P.). More recent expansions have been characterized 
as “culturally induced”—resulting from opportunistic 
individuals who have responded to situations created 
by human conditions such as the heaps of carrion beef 
that littered Arizona in 1891-1893 (Gehlbach 1981). 

In the U.S. and northern Mexico, coatis live 
mostly in the pine-oak and riparian woodlands of scat¬ 
tered mountain ranges, chiefly at elevations of 1,400 
to 2,000 m (4,600-6,500 feet), in a type of vegetation 
known as Madrean Evergreen Woodland (Brown 1973). 
They also range higher into coniferous forests and oc¬ 
casionally down into desert grassland and scrub. They 
can move easily across intermontane valleys through 
grassland and desert by following wooded riparian cor¬ 
ridors. They may even cross some treeless stretches 
in their northern push, but tree-sized vegetation is a 
requisite for resident populations (Gehlbach 1981). 

In contrast to most procyonids, coatis are mainly 
diurnal in activity and maintain a matriarchal social 
system with the females and young males in bands 
and the adult males usually solitary (Kaufmann 1962). 
Population densities are highly variable within years, 
but are lower in the southwestern U.S. than in the trop¬ 
ics, and there is some suggestion that at Nasua narica’s 
northern range limit the populations may be nomadic 
or migratory (Kaufmann etal. 1976). Like any species 
at the edge of their range, coati populations exhibit 
extraordinary flux, as natural and cultural factors exert 
unusually stressful limiting or powerful enhancing ef¬ 
fects. In 1960-1961, an epidemic of canine distemper 
reduced coati populations throughout Arizona, while 
a 32-inch snowfall in 1967-1968 drove the recover¬ 
ing Peloncillo-Guadalupe mountains population out 
of the southern part of that mountain range (Gehlbach 
1981). For these reasons, there has been concern about 
the conservation status of the species in the U.S. with 
suggestions that the species likely warrants complete 
legal protection. 


Marginal records for coatis in the U.S. gener¬ 
ally fall into two categories—occasional wanderers, 
and released or escaped captives. Most of the records 
are based only on sightings, and some of them are 
questionable. Most of the marginal records are of 
single animals. Coatis also commonly are acquired 
as pets because of their intelligence, inquisitiveness, 
and the charismatic appeal of the young. However, by 
their third year, coatis often become quarrelsome and 
destructive, and often are released back into the wild, 
where feral breeding populations may become estab¬ 
lished in more or less suitable habitat (see Kaufmann 
et al. 1976 for a thorough discussion). 

Herein we review the status of the coati in Texas 
with the following objectives: (1) to review and an¬ 
notate published historical records of the coati; (2) to 
present notes and other appropriate documentation 
concerning recent sightings of coatis made by profes¬ 
sional and amateur naturalists as assembled and sum¬ 
marized by the authors; (3) to assess these observations 
and records with regards to the natural history of the 
species at the northern limits of its range; and (4) to use 
this information base to make more accurate forecasts 
and assessments about the population and conservation 
status of N. narica in the state. 

Even though much of the information presented 
in this publication is not confirmed with “concrete” 
documentation, and especially in consideration of how 
little information is available about the coati in Texas, 
we believe the reported observations have value in 
suggesting areas of the state where vigilance should 
be exerted to determine if coati populations are present 
and/or possibly becoming established. Many natural 
history reports at the end of the 19th and beginning of 
the 20 th centuries made use of observations and records 
provided by local landowners or naturalists, and field 
agents were instructed to seek out such information 
(for example, see Bailey 1905 and other publications 
of the North America Fauna series published by the 
U.S. Biological Survey). This was deemed a neces¬ 
sary practice because specimen collecting was scat¬ 
tered and incomplete, especially for many of the more 
charismatic and rare species, and local residents were 
considered a valuable resource to fill-in distribution 
records when actual specimens could not be obtained. 
As we demonstrate, this situation is similar with regard 
to the coati in Texas. 


SCHMIDLY ET AL.-RECORDS AND STATUS OF NAS 11 A NARICA IN TEXAS 


129 


Historical Specimen and Sighting Records from the Literature 


There are 13 historical accounts in the literature 
representing 22 reports of coatis in Texas, but only 
three of these, representing four individuals, have been 
documented with scientific specimens. Most, but not 
all, of these records should be regarded as “confirmed” 
as discussed below. 

(1) The first published report of a coati in Texas, 
and the U. S., was reported in 1905 by Vernon Bailey 
who documented a specimen collected in 1877 at 
Brownsville, Cameron County, by J. C. Merrill. In a 
footnote to the species account, Bailey commented that 
this individual might have been an imported animal that 
escaped from captivity. This specimen is deposited in 
the mammal collection of the U.S. National Museum 
in Washington, D.C. 

(2) The next documented report of a Texas coati 
did not appear until 61 years later, when Taber (1940) 
reported a Mexican trapper catching a coati or “chulo” 
in July 1938, near Eagle Pass in Maverick County on 
the American side of the Rio Grande. With regard to 
the provenance of this record, Taber (1940) speculated 
“that the specimen probably was a pet that had escaped 
from captivity in Piedras Negras, just across the bor¬ 
der, to which place they often are brought for sale to 
American tourists.” A third record also was included in 
the report by Taber (1940). It was obtained 10 January 
1939, some 20 miles below Boquillas in the Dead Horse 
Mountains of Trans-Pecos Texas in what is likely now 
Big Bend National Park (BBNP). A game warden had 
mistaken the skin for an illegally taken river otter and 
confiscated it from a Mexican trapper before subse¬ 
quently presenting it to the Texas Cooperative Wildlife 
Collection (now Biodiversity Research and Teaching 
Collections) at Texas A&M University (catalogued as 
number 1412, skin & skull, prepared by R. L. Peterson 
and P. Goodrum, but missing since November 1966). 

(3) Afourth record was reported by Davis (1943): 
“In late February, 1943, Archie Kelly a resident of 
Concan, Texas, encountered a large coati, which his 
three collie dogs had treed, on the Rio Frio, about 40 
miles north of Uvalde, Uvalde County.” However, 40 
miles north of Uvalde is in Real and not Uvalde County 
and, therefore, we follow Goetze (1998; see below) in 


listing the record from Real County. The animal was 
an adult male measuring 142.2 cm (56 inches) in total 
length. Mr. Kelly was of the opinion that at least one 
other coati ranged in that locality, which led Davis 
(1943) to speculate that “this typically Mexican species 
is definitely extending its range into southern Texas and 
that it will likely become established.” When Taylor 
and Davis (1947) published the first edition of The 
Mammals of Texas , they described the distribution of 
the coati as “southern Texas from Brownsville to the 
Big Bend region of the Trans-Pecos.” 

(4) Another gap of almost two decades occurs 
before three additional records were reported by Na¬ 
tional Park Service personnel in BBNP—one from the 
U.S. side of the Rio Grande at Boquillas, a second from 
the Chisos Mountains in 1959, and the third from the 
Chisos Basin in 1966. These records were character¬ 
ized as representing “occasional pioneers from Mexico” 
(Kaufmann et al. 1976). 

(5) In the 1960 edition of The Mammals of 
Texas , Davis listed the coati as occurring in five Texas 
counties—Aransas, Brewster, Cameron, Maverick, 
and Uvalde counties. The basis for the Aransas county 
record was not explained in the species account, and the 
same five county records were listed in the 1966 edition 
of the book. It has been common practice in the various 
editions of The Mammals of Texas not to give specific 
details of records, many of which were provided to Da¬ 
vis and subsequent authors by game wardens and other 
personnel of the Texas Parks and Wildlife Department 
or by experienced local naturalists. 

(6) Halloran (1961) reported two sightings of 
coatis on the Aransas National Wildlife Refuge in 
Aransas County: “J. Clark Salyer II and former Refuge 
Manager James O. Stevensen saw a coati at Mustang 
Fake April 28, 1939. Another coati was seen on May 
15 of the same year.” It is possible these two reports 
represented the source of Davis’ 1960 inclusion of 
Aransas County as part of the coatis range in Texas. 

(7) The 1974 edition of The Mammals of Texas 
listed a record from Kerr County, again without in¬ 
cluding details, thus increasing the number of coati 


130 


Clyde Jones Memorial Volume 


county records to six. Similarly, the 1988 edition of 
this book listed the same six counties. To date there 
has not been any confirmation of the provenance for 
the Kerr County record. 

(8) In August 1975,1. Poglayen reported in a per¬ 
sonal communication to John Kaufmann a road-killed 
coati 50 kilometers (km) west of Abilene in Taylor 
County (Kaufmann et al. 1976). Given the distance 
from Abilene to the Mexican border (about 400 km or 
240 mi), the authors speculated that “the animal was 
probably an escaped captive.” 

(9) Another gap of almost two decades occurred 
before two records were documented and reported 
from Victoria County along the Texas coast (Henke 
and Young 1997). The first sighting occurred on 27 
July 1994, in riparian habitat approximately 1 mile 
southwest of the Guadalupe River and 2.7 miles east of 
U.S. Highway 77. The second sighting occurred on 29 
April 1995, when a coati was seen crossing State Road 
175 approximately 2 miles north of the intersection of 
Highway 77 and State Road 175 (now State Highway 
91). These sightings constituted the northernmost 
occurrence of coatis in the Gulf Prairies and Marshes 
vegetational area of Texas (Henke and Young 1997). 
The authors of this report offered the following inter¬ 
pretation of these records: “We believe it unlikely that 
the observed coatis were escaped pets or zoo specimens. 
The only zoo in South Texas that has coatis is the Texas 
Zoo, which is located in Victoria, Texas. They reported 
no missing specimens. Regional game officials were 
not aware of individuals keeping or breeding coatis 
locally.” Interestingly, these two records are from the 
same river-bottom habitat patch that is currently intact. 

(10) The 1994 edition of The Mammals of Texas 
by Davis and Schmidly listed nine counties with re¬ 
cords of coatis, including the previous six counties 
mentioned, plus new records of occurrence in Hidalgo, 
Starr, and Webb counties. Again, the details for the 
new records were not provided in the publication. The 
records for Webb County became available to Schmidly 
in 1982 when he examined two specimens collected in 
1901 at Laredo, Webb County, and housed in the mam¬ 
mal collection of the Philadelphia Academy of Natural 
Sciences. The two Laredo specimens (ANSP 6096 and 
6233) were donated to the collection by the Zoological 


Society of Philadelphia, which means they were zoo 
animals. ANSP 6096 is a skin with skull and skeleton 
but no sex indicated. External measurements are: total 
length, 42 inches (106.7 cm); tail 20.25 inches (51.4 
cm); and hind foot 3.5 inches (8.9 cm). ANSP 6233 is 
a male preserved as a skull and body mount. The skin 
tags for both read “Texas, Webb County, Laredo” with 
a date of “February 11, 1901.” These animals could 
have easily been pets purchased across the border in 
Nuevo Laredo, Mexico, for display at the zoo and then 
later donated to the collection. Two of us (Schmidly, 
DJS; and Karges, JK) recall from field-work in the 
borderlands not far from the U.S./Mexican border that 
roadside vendors commonly sold wild animals, includ¬ 
ing coatis, which potentially could constitute a source 
of transported and subsequently released pet animals. 

(11) Jones and Frey (2013 ), in their report on the 
mammals of Padre Island National Seashore (PAIS), 
described four reliable reports of coatis on the island: 
on the west side of the island adjacent to beach mile 
20, and at the entrance to PAIS in the early 1990s; a 
second report in January 1996 of an individual digging 
for crabs in the fore-dunes at beach mile 29; and photo¬ 
graphs of a group of coatis sighted in 2005 at the Best 
Western Hotel (14050 South Padre Island Drive) adja¬ 
cent to Packery Channel at the northern edge of North 
Padre Island (Nueces County, 4.62 kilometers south 
and 5.84 kilometers east of Flour Bluff). From these 
records the authors concluded, “although coatis are 
probably present within PAIS, their status is unknown.” 

(12) In the 2004 edition of The Mammals of 
Texas, Schmidly added a record from Real County and 
described the distribution of the coati in Texas as fol¬ 
lows: “they are only known from the southern part of 
the state, from Brownsville northwest to the Big Bend 
region of the Trans-Pecos and east to Kerr and Victoria 
counties.” The provenance of the record from Real 
County came from Goetze (1998). A careful review 
of his species account for the coati reveals the basis of 
the record to be Davis’ (1943) publication of the record 
from Uvalde County that now has been determined to 
have been taken in Real County (see above). 

(13) In the most recent edition of The Mammals 
of Texas, Schmidly and Bradley (2016) summarized 
the distribution of coatis in the state as follows: “His- 


SCHMIDLY ET AL.-DISTRIBUTION AND STATUS OF NASUA NARICA IN TEXAS 


131 


torically, they were known from the southern part of 
the state, from Brownsville northwest to the Big Bend 
region of the Trans-Pecos and east to Kerr and Victoria 
counties. Today they are known only from the Big Bend 
and Padre Island areas.” The distribution map for the 
species (p. 278) encompassed 12 counties: Brewster, 


Maverick, Kerr, Real, Uvalde, Webb, Victoria, Aransas, 
Starr, Hidalgo, Cameron, and Nueces. The record from 
near Abilene in Taylor County was not depicted on the 
map. From the records presented below in this paper, it 
is clear that the Lower Rio Grande Valley should have 
been included as a part of the coati’s range in Texas. 


Other Observations of Coatis in Texas 


We have compiled a number of recent reports 
of coati sightings in the Big Bend region, in the area 
between Big Bend and Del Rio, and from several loca¬ 
tions in the Lower Rio Grande Valley. These sightings 
constitute the basis for the information presented in 
this section of the paper along with some additional 
reports from personnel of the Texas Parks and Wild¬ 
life Department (TPWD) as well as other naturalists 
working along the Mexico-Texas boundary (see Ac¬ 
knowledgments). 

The provenance of the various sightings includes 
“confirmed” documented records and others that are 
more tenuous and lack “concrete” supporting infor¬ 
mation and therefore can only be considered as “anec¬ 
dotal” sightings or reports. Many of the “anecdotal” 
accounts come from second-hand reports of sightings 
made by amateurs without formal experience or training 
and lack physical evidence (specimens or photographs) 
or detailed, accurate descriptions. Confirmed sightings 
are well documented with specimens, photographs, or 
were made directly by professional naturalists-mam- 
malogists with experience observing coatis. 

Big Bend Basin and Big Bend National Park (BBNP) 

The observations described below were gathered 
by Rob Dean (RD) from the official park natural history 
field observation records that are maintained on-site at 
BBNP headquarters in Panther Junction. The forms 
available for this purpose are completed and submitted 
by individuals who believe they have seen an unusual 
animal worth documenting; the “record form” includes 
several categories of information—species observed, 
observer (with contact information), date and time of 
observation, weather, and space for comments about 
the description, behavior, or other pertinent information 
regarding the sighting. 


According to RD, the wildlife sighting forms are 
available at each visitor center in BBNP and are offered 
to individuals upon request. National Park Service 
(NPS) staff often will ask a visitor to fill out a form 
if the sighting is deemed significant. The completed 
form is then checked by staff and the reporting party is 
thanked for contributing to the park wildlife database. 
For rare or unusual sightings, NPS staff will “quiz” 
the reporting party to ensure as much of a degree of 
certainty as possible. The card is then photocopied and 
placed in a public binder at park headquarters. Both 
the original and the copy are incorporated into the per¬ 
manent files of the Park’s Science and Resource Man¬ 
agement Office. All visitor sightings are considered to 
be “anecdotal” unless verified by a photograph or by 
NPS confirmation; sightings by professionals, such as 
researchers or other experts in the field, are considered 
to be “confirmed” (personal communication, Raymond 
Skiles, Wildlife Biologist and Wilderness Coordinator 
at BBNP, to DJS). 

Rob Dean provided copies of the observation 
records for coatis to DJS, who studied the species 
observation descriptions and arranged them into three 
categories—none (no physical description of the ani¬ 
mal; simply stated “saw coati”); vague (some mention 
of defining features but not enough for complete con¬ 
firmation); and accurate (sufficient detail provided so 
there can be little doubt of identification). An example 
of an “accurate” description would be, “pointed face, 
white eye rings, white across nose, small ears, tail 
without evident rings; compared to photo of coati, no 
doubt of identity.” An example of a “vague” descrip¬ 
tion would be, “adult crossing road from west to east; 
walking very slow; tail dragging ground; healthy, full 
coat.” The breakdown of the 40 sightings according to 
these categories was as follows: “accurate” (8 sight¬ 
ings); “vague” (23 sightings); and “none” (9 sightings). 


132 


Clyde Jones Memorial Volume 


Thirty-five of the sightings came from within the 
boundaries of BBNP and five were from just outside 
the boundary of the park. The latter group included 
sightings from: U.S. Highway 385, just north of the 
park entrance (February 2009 and December 2011); 
north of Study Butte, State Highway 118 (no date); 10 
miles north of Terlingua, State Highway 118 (October 
1988); and Contrabando Canyon in Big Bend Ranch 
State Park. Of these only the latter record can be con¬ 
sidered as confirmed (see below). 

Sightings from inside the park included the fol¬ 
lowing (with dates): 

a. Vicinity of Rio Grande Village (7): January 2013; 
August 2008; December 2006; February 1987; 
September 1998; May 1978; August 1977. 

b. Vicinity of Panther Junction (7): August 2008; 
November 2002; January 2000; July 1998; April 
1987; January 1983; November 1978. 

c. The Basin (7): December 2000; November 1999; 
August 1998; March 1998; January 1998; June 
1994; September 1987. 

d. Boquillas Canyon (3): March 1986; November 
1977; November 1964. 

e. Santa Elena Canyon (2): January 2010; Novem¬ 
ber 1998. 

f. Castolon (2): March 2006; May 1988. 

g. Single locations: Sam Nail Ranch (February 
2009); Hot Springs (March 2002); Burro Mesa 
(February 1999); Lower Canyons (June 1998); 
Laguna Meadows (May 1989); Dugout Wells 
(September 2014). 

The sightings occurred over a 50-year period 
(1964-2014) with observations recorded in 20 of those 
years: 1964 (1 sighting); 1977 (2); 1978 (3); 1983 (1); 
1986 (1); 1987 (3); 1988 (2); 1989 (1); 1992 (1); 1994 
(1); 1998 (6); 1999 (3); 2000 (2); 2002 (2); 2006 (2); 
2008 (2); 2009 (2); 2010 (2); 2011 (1); 2013 (1); and 
2014 (1). More than 25% (11 of 40) of the sightings 
occurred during a three-year period from 1998 to 2000. 
There have been no sightings since 2014. Combining 
all of the records, sightings were made in every month 
of the year according to the following breakdown: 
November, 7; January, 6; March and September, 5; 


February, 4; June and December, 3 each; April and 
August, 2 each; and 1 each in May, July, October. 
Twenty-seven of the sightings (67%) were made during 
daylight hours, and all but two of these involved single, 
solitary individuals. There were two sightings of two 
animals made in April and February 1987, respectively, 
from Route 13,14 miles west of Panther Junction (at 2 
p.m.), and from the Rio Grande Village Nature Trail at 
8 a.m. A sighting made on 8 August 2008, in a tree at 
the Rio Grande Village Store, by Imre Karafiath (ex¬ 
perienced bird observer in the park) was “confirmed” 
by an excellent photograph (see Fig. 1). 

The coati is regarded as “an occasional migrant, 
not a resident, with credible reports in BBNP occurring 
on a less than annual basis” (RD and Raymond Skiles, 
personal communication). 

Sightings in the Trans-Pecos Outside of BBNP 

These sightings typically lack precise locality 
information and dates, much less any form of confir¬ 
mation such as photos, and therefore are considered 
anecdotal, even though for the most part they were 
made by individuals known to be reliable naturalists, 
unless noted otherwise. These sightings are reported 
as told to Karges (JK) or Schmidly (DJS), including 
discussion of the basis for species identification. 

Presidio County .—Details of a young coati seen 
in a roadside boulder field along the state highway adja¬ 
cent to Big Bend Ranch State Park (BBRSP) were pro¬ 
vided by Mark Lockwood, Natural Resource Specialist 
with TPWD, to Clyde Jones (Texas Tech University) 
when he was working on mammals in that area. Big 
Bend Ranch State Park staff have now confirmed the 
sighting with a photograph from their archives. The 
animal was seen 1 mi. east of Contrabando Canyon on 
State Highway 170 on 29 September 2000 (M. Lock- 
wood, TPWD, in litt.). This is the same sighting that 
was included in the official sighting records maintained 
at BBNP headquarters (see above). 

Michael Huston, a mammalogist in the Depart¬ 
ment of Biology at Texas State University who studies 
mammals in the Big Bend region, recently informed 
DJS (6 September 2016) of coati sightings made by a lo¬ 
cal resident who works for a bentonite mining company 


SCHMIDLY ET AL.-DISTRIBUTION AND STATUS OF NASUA NARICA IN TEXAS 


133 


Figure 1. A White-nosed Coati in a tree at the Rio Grande Village Store in Big Bend National Park, 8 August 2008. 
Photo courtesy of Imre Karafiath. 


in the Terlingua Creek drainage in the southernmost 
part of the county. According to the report, the man 
claims to have regularly seen two coatis over the past 
several years near Agua Fria, which is a spring-fed 
pool in the upper, generally dry reaches of Terlingua 
Creek. According to Huston, the man is known to be 
“very observant with birds and animals.” 

Lower Canyons of the Big Bend (Brewster, Terrell 
and Val Verde counties ).—Bill Russ, a Texas Parks and 
Wildlife Department district biologist who lived and 
worked in Terrell County in the early 1990s, reported 
coati troop sightings along the Rio Grande made from 
boating (rafting and canoe) trip participants, and from 
the descriptions provided both JK and Russ agreed 
these were coati sightings. There was no indication of 
how many sightings, how many animals, dates, or any 
other details or natural history specifics. 


Recently, we were informed of two additional 
sightings of coatis just below Boquillas Canyon. 
Bonnie McKinney (personal communication to DJS) 
reported a close-up sighting of a single individual under 
the cane of a sand bar along the river in 1985. In the 
fall of 2011, Marcos Paredes, then anNPS river ranger 
for BBNP, and a party of companions floating the river 
told JK of seeing multiple coatis on the Coahuilan side 
of the riverbank at International Water and Boundary 
Commission (IBWC) mile marker 791 at Rabbit Ears 
Canyon (Canon El Guerro). One was photographed and 
JK has obtained a copy of that image. We consider both 
of these records to represent “confirmed” sightings. 

Pecos County/Crockett County (near the Inter¬ 
state 10 bridge over the Pecos River). —In the late 
1990s, Dean Hendrickson, an ichthyologist at the Uni¬ 
versity of Texas, reported to JK that he saw three adult 


134 


Clyde Jones Memorial Volume 


coatis run across the interstate highway in the vicinity 
of the Pecos River bridge and valley near Sheffield. 
Hendrickson has years of field experience in Arizona 
and the Sierra Madre Occidental in adjacent Chihua¬ 
hua and Sonora, Mexico, with lots of familiarity and 
encounters with coatis. For this reason, we consider 
this to be a “confirmed” sighting. 

Crockett County .—Jim Mueller, then on the Biol¬ 
ogy Department faculty at Sul Ross State University 
and now the U.S. Fish and Wildlife Service (USFWS) 
Refuge Zone Biologist based at the Balcones Canyon- 
lands National Wildlife Refuge, told JK that Josh Avey, 
a Texas Tech graduate student studying mule deer on 
a hunting ranch east of Iraan in northwest Crockett 
County, sighted a troop of 12 to 15 coatis while con¬ 
ducting spotlight surveys on the ranch in 1999. The 
ranch in question had bank-side property with the Pecos 
River which would lend some credence to a possible 
corridor for movement up the river. 

Terrell County .—At Cedar Station, between 
Langtry and Dry den, two teenage sons of Pat and Glenn 
Merkord (friends of JK) reported seeing two coatis 
feeding at daybreak as they watched from a deer stand 
near a baited corn feeder. The animals were feeding as 
it got light enough to see. According to JK, the whole 
family, including the sons, would have been able to 
discern between coatis and raccoons, gray foxes, or 
ringtails but we still consider the record “anecdotal.” 

Davis Mountains, Jeff Davis County .—Chris 
Durden (a lepidopterist then in the Biology Depart¬ 
ment at the University of Texas) told JK that he saw 
a coati in the upper Davis Mountains Resort region 
(east of Mt. Livermore) of the Davis Mountains on 21 
October 1974. In April 2012, Louis Harveson from 
the Borderlands Research Institute at Sul Ross State 
University, as part of a study of mountain lions in the 
mountains, reported that a coati was found attacked 
by a lion or coyote after having been caught in one 
of their traps. According to Jonah Evans (personal 
communication to DJS), the identification was based 
on inspecting the fur at an old lion kill site. Recently, 
on 11-12 June 2016, Chris Mallery, a herpetologist 
from the University of North Texas camping in Davis 
Mountains State Park, reported to JK sighting an animal 
matching the description of a coati on State Highway 


118 just outside of the park. F inally, Bryan Hughes, a 
biologist from Arizona familiar with coatis, informed 
DJS that he observed a coati on 19 August 2012 just 
before dark (7:30 pm) on Ranch-to-Market road 1832, 
in the northern part of the Davis Mountains. With the 
exception of the latter record, we do not regard any of 
the records as “confirmed.” 

Val Verde County .—Multiple anecdotal sightings, 
covering drainages of the Rio Grande, Pecos and Devils 
rivers, have been reported to JK from this county. In 
July 2014, a naturalist’s sighting came to JK’s attention 
of a reported solitary coati, followed a few months later 
by a second sighting from a ranch near Pumpville on 
tributaries of the Pecos River. The stewards for The 
Nature Conservancy (Jim and Bea Harrison) at the 
Dolan Falls Preserve provided documentation that some 
members of the Texas Ornithological Society’s board, 
while attending a meeting/retreat at the preserve in the 
late 1990s, saw a coati crossing the road in daylight 
hours just as they were departing the Devils River 
State Natural Area. When JK mentioned the sighting 
to Jim Finegan, former TPWD state natural area staffer 
and descendent of the original landowner family, he 
too mentioned casually that he had seen a coati on the 
natural area but without date or details. Joe Joplin, 
current TPWD Superintendent of the Devils River State 
Natural Area, reported that another TPWD employee 
familiar with coatis from previous Arizona experience 
told him about sighting a juvenile coati crossing Dolan 
Creek Road near the old Fawcett/Finegan Ranch head¬ 
quarters at 6:55 a.m. on21 April 2015. Rob Klockman, 
a graduate of the Department of Wildlife and Fisheries 
Sciences at Texas A&M University, reported two ad¬ 
ditional sightings from the county—in May 2005, from 
Loma Alta on U.S. Highway 277; and in May 2010 on 
U. S. Highway 90 just west of Langtry. F inally, Nathan 
Wells, another naturalist friend of JK, reported sighting 
a juvenile coati on 4 July 2015 near a road-cut along 
U.S. Highway 277 north of Del Rio. 

Observations in South Texas and the Lower Rio 
Grande Valley 

Fewer sightings have been reported in southern 
Texas than in the Big Bend and regions to the east along 
the border with Mexico, but the number of sightings has 
been increasing recently, including a recent capture of 


SCHMIDLY ET AL.-DISTRIBUTION AND STATUS OF NASUA NARICA IN TEXAS 


135 


a coati and its subsequent release by TPWD officials. 
These reports are documented as communicated by 
various individuals to JK or DJS and unless noted 
otherwise are regarded to be anecdotal sightings. 

Cameron County. —Tony Henehan, currently 
a TPWD biologist in Weslaco, reported to JK that a 
former TPWD biologist told him of sighting a coati 
crossing a road within the Las Palomas Wildlife Man¬ 
agement Area along the Arroyo Colorado at the Arroyo 
Colorado Unit north of Rio Hondo. No dates or other 
details concerning the sighting were recorded. 

Hidalgo County. —Javier de Leon, TPWD Super¬ 
intendent of the Estero Llano Grande State Park, told 
JK that in 2011 or 2012 (he could not be sure of the 
specific year) a maintenance worker reported 4-5 coatis 
on TPWD property a few miles east of the state park. 
The employee, who the Superintendent claimed knew 
the difference between coatis, raccoons, or opossums, 
saw the animals among tepeguaje and anacua trees 
within the IBWC floodway. Also, in 2012, Javier de 
Leon reported that a Bensen-Rio Grande Valley State 
Park volunteer told him that maintenance workers at 
Benson Grove RV Park (approximately 3 mi. directly 
north of the state park) captured a coati while trying to 
trap nuisance opossums, describing the animal as pos¬ 
sessing “a long nose and long brown tail with rings.” 
That animal was released unbeknownst to park staff 
at Bentsen-Rio Grande Valley State Park, southwest 
of Mission (de Leon in litt.). 

Jonah Evans, the state mammalogist with TPWD, 
told DJS about a coati that was captured in a residential 
backyard in Alamo, Hidalgo County, on 17 April 2016 
by municipal animal control personnel. The animal was 
brought to the Gladys Porter Zoo in Brownsville where 
it lived for some time before TPWD personnel became 
aware of the circumstances. TPWD officials later re¬ 
leased the animal at the Anacua Unit of Las Palomas 
Wildlife Management Area near Santa Maria, Cameron 
County, on 26 July 2016. This record is regarded as a 
“confirmed” sighting. 


Maverick County. —Kathy Pine, at the time a 
staff member of The Nature Conservancy in Houston, 
told JK in the early 2000s that while spending time on 
a ranch in this county, she saw of troop of coatis. She 
was emphatic they were coatis. 

McMullen County (Tilden). —In July 1994, Rob 
Klockman (see above) reported seeing a coati at Tilden, 
which is the county seat and lies at the intersection of 
State Highways 16 and 72 in the north-central part of 
the county. 

Western Willacy County/Eastern Hidalgo Coun¬ 
ty. —Tony Henehan, a TPWD biologist responsible for 
Wildlife Management Areas, reported that a scorched/ 
road-killed coati carcass was obtained following “the 
burning of a sugar cane field in Willacy County a couple 
of miles north of La Villa, Texas.” The specimen was 
salvaged and will be deposited in an academic museum 
research collection. The animal was reported as coming 
from Willacy County, although La Villa is in eastern 
Hidalgo County near the county line. Although this is 
considered by us to represent a “confirmed” record, the 
wildlife technician who assisted with the recovery of 
the specimen unfortunately could not recall the exact 
year of the discovery, only that it was in either the 
month of August or September. 

Webb County. —Jim Goetze, a mammalogist in 
the Science Department at Laredo Community Col¬ 
lege, reported to DJS (9 September 2016) that a live 
coati is currently on display at the Lamar Bruni Ver¬ 
gara Environmental Science Center on the campus at 
the community college. According to the report, the 
animal (a male) was procured from a family who had 
imported and raised it for at least two years. Also, Goe¬ 
tze reported to DJS that he receives occasional reports 
of coati sightings from U. S. Border Patrol personnel 
and hunters, but that in 21 years of biological work in 
Webb County and the surrounding area he has never 
seen a coati. 


136 


Clyde Jones Memorial Volume 


Surveys Conducted by Professional Mammalogists within the Range of Coatis 


Several long-term field surveys of mammals 
have been conducted by professional mammalogists 
working in the geographic regions where many of the 
aforementioned sightings were made. These studies 
involved extensive trapping (cage traps and in some 
instances camera traps) for carnivores as well as day¬ 
light and nocturnal excursions to search for mammals. 
These are listed here along with appropriate comments 
regarding the presence of coatis in the survey areas. 
The results of these studies have been reported in sci¬ 
entific publications, agency reports, or made as personal 
communications to DJS. 

Big Bend National Park (1974-1976). —David 
Schmidly and Robert Ditton (1976), along with their 
graduate students, spent two years assessing riparian 
sites in BBNR They visited 64 separate sites along 
the river between the park boundaries, traveling by 
vehicle along the River Road and spent dozens of days 
and nights “river-rafting” in Santa Elena, Mariscal, and 
Boquillas canyons. Their efforts included trapping 
for both small and large mammals as well as day-light 
and night-time observations for large mammals. They 
never saw a coati or received a report of one in the 
park during this time frame. Both Rio Grande Village 
and Santa Elena Canyon, where coati sightings were 
subsequently made, were visited and sampled by them 
on multiple occasions. 

Harte Ranch, Big Bend National Park, Brewster 
County (1991-1993). —Clyde Jones, Frank Yancey, and 
Richard Manning conducted an extensive field survey 
of mammals in this newly acquired parcel (57,000 
acres) along the northern boundary of the Park. Despite 
considerable effort to capture and observe carnivores, 
they never captured, sighted nor received any reports of 
coatis in the area (Jones et al. 1993; Yancey et al. 2006). 

Big Bend Ranch State Park, BBRSP (1994- 
1995). —This large state park (one of the largest state- 
managed land areas in North America) is located in 
southeastern Presidio County just west of BBNP. Frank 
Yancey and Clyde Jones collected and observed mam¬ 
mals at more than 300 localities throughout the state 
park. The published report (see Yancey 1997) included 
this statement about coatis: “Local residents have 


reported the occurrence of N. narica in BBRSP, but 
these reports are unsubstantiated. The species prefers 
woodlands and rocky areas... both present in BBRSP, 
so an occasional individual may wander into the park 
from Mexico.” The incidental observation of a coati 
was confirmed by a photograph taken after the Yancey 
publication (discussed earlier). 

Chinati Mountains State Natural Area, Presidio 
County (2002-2010). —Clyde Jones, his students, and 
colleagues surveyed mammals in this area over a period 
of almost a decade. Located just up river and northwest 
of BBRSP, they collected throughout the mountains, 
but found no evidence of coatis and made no mention 
of them in their published report (Jones et al. 2011). 

Davis Mountains, Jeff Davis County (1998- 
2002). —For four years, Robert DeBaca (2008) sur¬ 
veyed mammals in the Davis Mountains, including 
The Nature Conservancy’s (TNC) Davis Mountains 
Preserve, Davis Mountains State Park, Balmorhea State 
Park as well as Phantom Spring and Sandia Springs 
Preserve. He also examined museum and literature 
records and found no reports of coatis in the area. As 
explained above, there have been a few unconfirmed 
sightings of coatis in the Davis Mountains proper since 
the study by DeBaca. A camera trap study on the TNC’s 
Davis Mountain Preserve by staff of the Borderlands 
Research Institute at Sul Ross State University appar¬ 
ently has not produced any sightings of coati. 

Indio Mountains Research Station, Hudspeth 
County (1993-present). —The University of Texas at El 
Paso (UTEP) has maintained a field station since 1993 
in these mountains, which are about 40 miles southwest 
of Van Horn. Jerry Johnson, Professor of Biology at 
UTEP and a long-time colleague of DJS, provided 
these comments to DJS about coatis in the region: “I 
have been actively engaged in field work in that area 
since 1973 during both daylight and nighttime hours, 
and have never seen a live or road-killed coati, nor has 
any local ever mentioned seeing them around here”. 

Amis tad National Recreation Area, Val Verde 
County (1976-1977; 2006-2007). — Several extensive 
surveys of mammals in this area did not produce any 


SCHMIDLY ET AL.-DISTRIBUTION AND STATUS OF NASUA NARICA IN TEXAS 


137 


records or sightings of coatis. Robert Ditton and DJS 
(1977) surveyed selected sites with high human activ¬ 
ity, collecting both small and large mammals as well 
as conducting daytime and nighttime observational 
surveys. J. M. Mueller and his students at Sul Ross 
State University conducted an extensive trapping sur¬ 
vey of mammals in the Recreation Area in 2006-2007, 
and they did not report any sign of coatis in the area 
(Bahm et al. 2008). 

Devil’s River State Natural Area (2001, north 
unit). — Brant and Dowler (2001) surveyed mammals, 


including mesocarnivores, at the Natural Area for 
several years and did not sight or report any records 
of coatis. 

Edwards Plateau (1989-1994). —Jim Goetze 
(1998) actively collected and searched for mammals 
on the Edwards Plateau over a 5-year period and never 
recorded any sightings of coati in the area. His pub¬ 
lished report included a reference to the record from 
the border of Uvalde and Real County published by W. 
B. Davis in 1943 (see above). 


Camera Trap Surveys 


In recent years, camera traps have become an 
important tool in wildlife research and management, 
especially for surveying carnivore species (Foster and 
Harmsen 2012). Camera traps provide tools to more 
thoroughly survey species over a larger area than may 
be possible with other survey techniques, particularly 
in remote areas with rough terrain (Silveira et al. 2003). 
We have become aware of a number of camera trap sur¬ 
veys in areas where coati sightings have been reported 
and provide details below. 

Dennison et al. (2016) placed paired trail cam¬ 
eras at 38 locations throughout the Davis Mountains 
on TNC’s Davis Mountain Preserve and two adjacent 
private ranches in Jeff Davis County. The habitat is 
very similar to that preferred by coatis at the northern 
edge of their range (montane evergreen forests, wood¬ 
lands and savannah, and riparian gallery woodlands). 
Cameras were activated at each site for a minimum 
of three months between June 2012 and March 2013. 
Feral hogs ( Sus scrofa) and gray fox (Urocyon cine- 
reoargenteus ) were the most widespread species, each 
observed at 33 of 38 camera locations. Mountain lions 
(Puma concolor ) were observed at 22 of the 38 camera 
locations. Mesocarnivores recorded included coyote 
(Canis latrans ), skunk (no species designation), bobcat 
(Lynx rufus ), raccoon (Procyon lotor ), and ringtail (Bas- 
sariscus astutus). There were no camera observations 
of coatis. 

James Eddy (personal communication to DJS), a 
graduate research assistant in the Borderlands Research 


Institute at Sul Ross State University, placed 10 cameras 
over about 600 acres at Elephant Mountain Wildlife 
Management area from April 2015 until August 2016. 
This area is about 26 miles south of Alpine in Brewster 
County. The objective of the project was to study quail, 
but several mesocarnivores, including badger (Taxidea 
taxus ), black bear (Ursus americanus), bobcat, coyote, 
gray fox, raccoon, and skunk (no species given), were 
detected but no coatis. 

Raymond Willis (personal communication with 
DJS), Director of the Dalquest Research Station of 
Midwestern State University, and some of his students 
have placed 20 camera traps on Terlingua Ranch in 
southern Presidio and Brewster counties since 2013. 
This 3,000-acre property is located on the northeast 
border of BBRSP about 20 miles from the Mexican 
border. Their cameras have produced regular sight¬ 
ings of mountain lion, coyote, raccoon, bobcat, gray 
fox, a few ringtails, and one or two black bear but no 
sightings of coati. 

Michael Huston (personal communication with 
DJS) and his students in the Wildlife Biology program 
at Texas State University have had an array of 8 camera 
traps in the canyons on the west side of the Christmas 
Mountains just north of BBNP since February of 2015. 
Their traps also produced no sightings of coatis. 

In 2014 and 2015, Skyler Stevens (2016), a gradu¬ 
ate student in the Department of Natural Resources 
Management at Sul Ross University, conducted an 


138 


Clyde Jones Memorial Volume 


extensive camera trap study in BBNP. Fifty-eight 
cameras were deployed over a 450 square kilometer 
grid covering the Chisos Mountains, parts of the Sierra 
Quemada, Burro Mesa, and some of the flats north and 
east of the Chisos. The purpose of the project was to 
document occurrences of mountain lions and their 
prey. The cameras were placed in areas expected to 
capture animal movement such as washes, saddles in 
ridges, canyons, and at convergences of game trails. 
Over 14,000 trap nights produced 500,000 pictures that 
were sorted and analyzed. The results produced more 
captures of mesocamivores than big cats; among the 
procyonids, a few raccoons and several ringtails were 
noted but coatis were never observed, not even a suspi¬ 
cious photograph. In addition to the Sul Ross project, 
BBNP has maintained six cameras in diverse habitats 
throughout the part for six years with no sightings of 
coatis (Raymond Skiles personal communication to 
DJS). 

Marc Cancellare (personal communication to 
DJS), a graduate student with Richard Kazmaier at 
West Texas A&M University, placed 16 cameras at 
Black Gap Wildlife Management Area in southeastern 
Brewster County from September 2014 to October 2015 
(total of 6003 camera days and 360,934 images) and, 
to date, has not recorded any photos of coatis. Over 
the same time-frame, camera traps also were placed 
on the privately-owned Buckhollow Ranch (Real and 
Uvalde counties) and TNC’s Independence Creek Pre¬ 
serve (Terrell County) with no reported observations 
of coatis at either site. 

Dowler et al. (2016) made extensive use of 
camera traps to sample mesocamivores on the recently 


acquired Dan A. Hughes Unit at the Devil’s River State 
Natural Area, located 35-45 km north of Del Rio in 
Val Verde County, between February 2013 and August 
2015. Twenty-two camera traps were placed along 
dry washes or in areas where animals were thought to 
be passing between February 2013 and August 2015 
(total of 3,547 camera days). Also, camera traps were 
accompanied by 996 cage-trap nights using Tomahawk 
Live Traps. Mammals recorded by the camera traps 
included: opossum ( Didelphis virginiana), jackrabbit 
(Lepus californicus), porcupine ( Erethizon dorsatum), 
gray fox, bobcat, mountain lion, hog-nosed skunk 
(Conepatus leuconotus ), ringtail, and raccoon. There 
were no captures or photographs of coatis made during 
this period even though previous anecdotal sightings 
of coatis existed for this area (see above). 

Mike Tewes (personal communication to DJS), 
a Research Professor at the Caesar Kleberg Wildlife 
Research Institute at Texas A&M Kingsville, has 
conducted extensive camera trapping and observa¬ 
tion studies for ocelots ( Felis pardalis) in the South 
Texas brushlands and Lower Rio Grande Valley since 
1985. He has never encountered a coati “after prob¬ 
ably 50,000 cage-trap nights and over 50,000 camera 
nights in south Texas.” He went on to comment that 
he has trapped and collared coatis in northeast Mexico 
as well as photographed them in different locations. 
Elaborating further, Tewes told DJS he talked with Mr. 
Jimmy McAllen, who owns several large ranches just 
north of the Rio Grande Valley, and who keeps track 
of wildlife oddities or unusual sightings. McAllen said 
he had never found a coati, but that he did know of one 
person who had released a captive animal. 


Discussion and Recommendations 


What can be made of all of this—a handful of 
published historical records over a 139 year period, 
several of which appear to represent “escaped pets,” 
with a few recent confirmed sightings; numerous “an¬ 
ecdotal” sightings over decades from scattered areas 
along the Texas border with Mexico but mostly adja¬ 
cent to the Rio Grande and Devils River and recently 
from the Lower Rio Grande Valley; the absence of any 
coati sightings during extensive field collecting surveys 


made over the past 40 years by professional mammalo- 
gists; and a complete lack of coati photos from seven 
camera trap studies conducted over the past five years 
in areas where coatis have been incidentally sighted 
during that time? 

Although much of the documentation assembled 
and discussed herein is “anecdotal” and not accompa¬ 
nied by “hard evidence” of occurrence, it does provide a 


SCHMIDLY ET AL.-DISTRIBUTION AND STATUS OF NASUA NARICA IN TEXAS 


139 


basis for some useful interpretations that can be refined 
over time as more detailed and accurate information 
becomes available. Some studies advocate use of 
anecdotal data, whereas others demand more stringent 
evidentiary standards such as only accepting records 
verified by physical evidence, at least for rare or elusive 
species. Frey et al. (2013) demonstrated that occur¬ 
rence datasets based on anecdotal records can be useful 
when inferring species distributions, provided that data 
are used only for easily-identifiable species and are 
based on robust modeling methods. In the American 
Southwest (New Mexico and Arizona), they were able 
to demonstrate that the predicted distribution of the 
coati based on anecdotal occurrence records was simi¬ 
lar to datasets that only included physical occurrence. 

Coatis are highly distinctive in appearance and 
behavior and are unlikely to be misidentified by careful 
or knowledgeable observers. Because coatis are diurnal 
and mostly active during the day, anecdotal reports are 
more likely to represent accurate identifications than 
would be expected for carnivore species that typically 
are nocturnal (e.g., opossums, raccoons, and ring¬ 
tails). In addition, because coatis are an unusual and 
relatively rare species in Texas, encounters are likely 
to be remembered. For these reasons, we propose that 
many sightings of coatis in Texas, as documented in 
this paper, represent valid observations and provide 
useful information. Although some of these observa¬ 
tion records may be considered doubtful, others are 
definitely reliable and are too numerous to ignore. 
Some of the included sight records undoubtedly repre¬ 
sent exceptional wanderings by wild individuals, most 
likely adult males, but many of these animals appear 
to be released or escaped captives. 

Fortunately, there is a good knowledge of the 
natural history of the coati at the northern margin of 
its range in Arizona where there are resident, breeding 
populations (see Kaufmann et al. 1976 for details). 
This provides a useful background for interpreting the 
status of the species in Texas. 

Observations of coatis in Texas are scattered 
over time and highly periodic. Gaps in observations, 
representing spans of several years and even decades, 
are evident. Records, represented by specimens, pho¬ 
tographs, documented sightings, and recent anecdotal 


sightings included in this paper, are now available from 
19 counties in the state including 13 counties with 
“confirmed” records (see Fig. 2). With a few excep¬ 
tions, most of the sightings have been made along the 
Texas-Mexico borderlands (within 100 miles of the 
border) from Big Bend east to Del Rio and south to 
Brownsville. 

Most of the documented sightings in Texas have 
been of single animals. Sightings of multiple individu¬ 
als or troops are among the most poorly documented 
of the records. Such a pattern would suggest the 
observations primarily represent marginal records of 
occasional wanderers and released or escaped pets. 
Of the scattered records from Texas, only those from 
the Big Bend’s Rio Grande area and perhaps from 
the Devils River basin likely represent true wander¬ 
ers from Mexico. These two rivers, together with the 
Pecos River, represent possible dispersal corridors for 
wandering coatis from Mexico to make their way fur¬ 
ther inland as they are known to use river and stream 
corridors as well as springs. 

Unlike Arizona, where coatis have been resident 
year round for over a century and exist as breeding 
populations, there is no evidence that a breeding popu¬ 
lation of coatis has been established or exists in Texas. 
While young have been sighted, they do not appear in 
troops as would be expected if a breeding population 
had been established—coatis breed annually so mixed 
age populations would be expected. Hundreds of troop 
sightings, including adult females with young males 
and females, are available from Arizona. In Arizona 
most mating apparently takes place in April, the bands 
break up before the young are born in June, and the fe¬ 
males with their new litters re-gather with the yearlings 
of both sexes in August. 

Our observations are consistent with the interpre¬ 
tation of Fred Gehlbach (1981), who spent several years 
observing coatis along the U.S.-Mexico Borderlands, 
and concluded “all of the Texas records represented 
solitary, wandering males or escaped and released 
pets and that there were no family bands of this spe¬ 
cies within a hundred miles of the Texas border.” This 
interpretation is reinforced by the fact that numerous 
long-term surveys by professional mammalogists, as 
well as several recently conducted camera trap studies, 


140 


Clyde Jones Memorial Volume 


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Figure 2. County records of coatis in Texas. Closed circles represent counties with “confirmed records”; open circles 
represent counties with only “anecdotal” sightings; open circles followed by a question-mark are from counties with 
questionable documentation; half-filled circles represent counties with both “confirmed” and “anecdotal” records. 


in the areas where many of the confirmed/anecdotal 
sightings have been reported failed to document any 
evidence of established coati populations. Perhaps 
in some of the areas where clusters of sightings have 
been made over decades, such as BBNP, rare popula¬ 
tions have temporarily flourished beyond their normal 
breeding range but did not persist in marginal habitat 
and likely were killed off by periods of drought and 
cold that eliminated them in relatively short intervals. 


Surveys from southeastern Arizona and south¬ 
western New Mexico show that the distribution of 
coatis corresponds almost exactly to that of the Encinal 
and Mexican Oak-Pine Woodland as mapped by Brown 
(1973). They also are known from lower elevations 
in cottonwood-sycamore-willow associations near 
streams and springs, and marginal records have been 
recorded in Chihuahuan Desert scrub and grassland 
habitat and in riparian areas surrounded by desert. 


SCHMIDLY ET AL.-DISTRIBUTION AND STATUS OF NASUA NARICA IN TEXAS 


141 


This pattern matches very well with the observations 
from BBNP where the most numerous sightings were 
in Rio Grande Village (cottonwood-willow), the Basin 
(pinyon-oak-juniper), and the vicinity of Panther Junc¬ 
tion (desert scrub habitat). 

Several of the sightings in BBNP were from 
places popular with campground campers along the 
River Road adjacent to the Rio Grande (e.g., Rio 
Grande Village, Santa Elena Canyon, Castolon, and Hot 
Springs). Gehlbach (1981) noted that solitary coatis 
are prone to beg marshmallows and raid garbage cans 
at night. Also, at such places he noted they often shift 
from diurnal to nocturnal habits in apparent response 
to nocturnally unattended trash cans and diurnal camp¬ 
ground hassle. He reported at Chiricahua National 
Monument seeing a coati “table” seven campers in five 
seconds one evening. 

The source population for Texas coatis is most 
likely the mountains along the northern border of the 
Mexican states of Coahuila, Nuevo Leon, and Tam- 
aulipas. Coatis have not been documented east of the 
Rio Grande in New Mexico nor or there any observa¬ 
tions along the Texas border from El Paso to Presidio 
making it highly unlikely that animals from Arizona 
and western Chihuahua, Mexico, would ever disperse 
into Texas. 

Baker (1956) in his study of Coahuila, Mexico, 
mammals wrote that he had “received no definite re¬ 
cords of the coati along the Rio Grande in Coahuila.” 
However, McKinney and Villalobos (2004) and McK¬ 
inney (2012), who have conducted wildlife studies for 
almost 20 years in the Maderas del Carmen/Sierra del 
Carmen mountains of northern Coahuila on the border 
with BBNP, are confident that the coati has become 
established there through recent range expansions and 
is now a permanent resident. McKinney and her associ¬ 
ates have made numerous sightings of coatis, including 
several troops in the pinon-juniper, pine-oak, and fir 
woodlands from 4,500 to 6,000 feet, always in or near 
riparian areas, going back to 2002 and 2003. They 
consider the coati to be a rare resident of the moun¬ 
tains; significantly all of their sightings were of groups 
and not solitary individuals. According to McKinney 
(personal communication to DJS): “I am 100 percent 
confident this is a resident breeding population that is 
not large but widely scattered over the landscape,... 


and I suspect that the coati people have seen in the Big 
Bend area are for sure from the Carmens.” 

The Rio Grande, although probably a barrier to 
small mammals where it flows through deep canyons 
along part of the northern boundary of Coahuila, seems 
not to bar the passage of most mammals where the 
river’s banks are low (Baker 1956; Schmidly 1977). 
We have confirmed sightings from both sides of the 
Rio Grande in the Boquillas region of BBNP clearly 
supporting the interpretation that coatis are able to 
cross the river. 

In Chihuahua, coatis are known only from the 
western Sierras where they undoubtedly are a source 
population for Arizona and New Mexico, but they 
have not been recorded from the mountains nearest to 
the border of Texas (Anderson 1972). Similarly, the 
nearest mountains to the Pecos and Devils rivers, east 
of the Big Bend area, with possible coati habitat are the 
Serranias del Burro in northern Coahuila. Coatis have 
not been recorded there although no extensive mammal 
survey has ever been attempted from that mountain 
range which represents the northernmost sky island 
of the Sierra Madre Oriental in the state of Coahuila, 
Mexico. The area does include habitat similar to that 
in the Sierra del Carmen/Maderas del Carmen where 
coatis are known to occur. 

Source populations for coati in south Texas and 
the lower Rio Grande Valley would likely come from 
isolated mountain ranges near the border in the Mexican 
states of Nuevo Leon or Tamaulipas. In Tamaulipas, 
just below Brownsville, coatis have been recorded from 
the San Carlos Mountains, about 200 km (120 miles) 
south of the Texas border (Schmidly and Hendricks 
1984), and these mountains could be a potential source 
for animals to wander north and enter the lower Rio 
Grande Valley. But, an even more likely source popu¬ 
lation would be the Sierra Picachos between Roma, 
Texas, and Monterrey, Nuevo Leon. A Mexican biolo¬ 
gist and colleague of JK, Enrique Guadarrama, who has 
been studying black bears in these mountains, reports 
that coatis are common there which would make those 
mountains a much more likely proximal source of 
wandering animals (single males or troops) to enter the 
western Lower Rio Grande Valley. The Sierra Picachos 
are only 90 km (56 miles) from Roma, Texas. 


142 


Clyde Jones Memorial Volume 


The conservation status of the coati in Texas 
remains enigmatic. The species was listed as state 
endangered in Texas in 1993 but has since been down¬ 
graded to threatened by TPWD and ranked as an “S2” 
species. Such a ranking is used to designate imperiled 
populations at high risk of extinction, or elimination 
due to very restricted range, very few populations, steep 
declines, or other factors. The coati is listed as a Spe¬ 
cies of Greatest Conservation Need (SGCN) in Texas 
according to the Texas Conservation Plan (TCAP) 
[http://tpwd.texas.gov/land/tcap/sgcn.phtml] for the 
Chihuahuan Desert, Arizona-New Mexico Mountains, 
Edwards Plateau, and South Texas Plains ecoregions, 
and therefore warrants conservation attention and 
additional information on status and distribution in 
those areas. 

Coatis would be seriously impacted by degrada¬ 
tion of riparian woodland habitat in these areas because 
they require a sizeable area of habitat to maintain 
a viable population (Schmidly 2002). Given their 
tenuous status, John H. Kaufmann, who at one time 
was the coordinator of the Coati Study Project in the 
United States, recommended complete legal protection 
for these animals although no official listing was ever 
made and the species is not currently included on the 
endangered species list by the USFWS (Kaufmann 
1987). In New Mexico, the coati is an endangered spe¬ 
cies, under legal protection, and it may not be hunted 
or trapped. In Arizona, it is considered a nongame 
mammal and may be taken during an open season, with 
a bag limit of one per calendar year. In Mexico it is 
not considered endangered and there is practically no 
information on the status of its populations even where 
it is abundant (Ceballos 2014). In northern Mexico, 
hunting apparently has caused significant reductions in 
their populations (Gompper 1995). 

Additional study and information will be required 
in order to better predict the conservation status and 
long-term viability of the species in Texas. A more for¬ 
mal system for documenting and following up on veri¬ 
fied sightings should be developed and implemented. 
Observers should be encouraged to document incidents 
with photography and salvaged specimens (for road- 
killed animals) with accurate localities and circum¬ 
stances of the sightings. The same system should be 
put in place for observers utilizing camera traps. State 


agencies (TPWD, State Land Office), NGOs (The 
Nature Conservancy and Audubon Society), academic 
institutions (perhaps through the Texas Mammal Soci¬ 
ety) and interested landowners could cooperate in such 
efforts. If a troop of coatis is documented, professional, 
experienced naturalists should be funded to conduct 
ecological and behavioral studies that might include 
radio-collaring of individuals, obtaining tissue samples 
through non-lethal means to conduct genetic assess¬ 
ments of population structure and taxonomic affinity, 
and gathering other basic natural history information 
(food habits, reproductive patterns, movements, etc.). 

The success of the El Carmen—Big Bend Con¬ 
servation Corridor project (see McKinney 2012) offers 
much hope for the eventual establishment of a perma¬ 
nent, breeding coati population in Texas. This program 
is a cooperative effort by the U. S. and Mexican govern¬ 
ments, private conservation groups and area ranchers to 
provide and protect a corridor on both sides of the Rio 
Grande in the Big Bend region that will allow wildlife 
to move freely within an intact ecosystem (McKinney 
2006). Currently, the corridor includes over 400,000 
acres of land in the Sierra del Carmen and Maderas 
del Carmen and 47,000 acres of “wilderness” with no 
development at all along the Rio Grande. The corridor 
from Mexico literally comes across the border at the 
eastern end of Boquillas Canyon and follows the big 
valley to the north. The American black bear ( Ursus 
americanus) population has increased substantially 
on both sides of the border as a result of this habitat 
protection plan, and presently efforts are underway to 
re-establish bighorn sheep (Ovis canadensis ), mule deer 
(Odocoileus hemionus ), and pronghorn ( Antilocapra 
americana ). According to McKinney (email commu¬ 
nication to DJS) the coati is also considered a critical 
species in their project. This effort could eventually 
result in the establishment of a viable, sustainable 
coati population in the Chisos Mountains of the Big 
Bend region. 

Coatis wandering north along regions to the east 
of the Big Bend and the lower Rio Grande Valley would 
have to cross a broad area of desert-scrub habitat to 
become established in Texas, and while a few of them 
may continue to wander into this region along riparian 
woodland corridors they probably will not be able to 
permanently occupy such marginal habitat. However, 


SCHMIDLY ET AL.-DISTRIBUTION AND STATUS OF NASUA NARICA IN TEXAS 


143 


there could be some corridor potential in the Lower Rio 
Grande Valley, where recent and numerous sightings 
have been reported, from adjacent source populations 
in Tamaulipas and Nuevo Leon in Mexico and the 
units of protected TPWD and USFWS lands, as well as 
Audubon Texas’ Sabal Palm Sanctuary and The Nature 
Conservancy’s Southmost Preserve. 

Finally, there is one other aspect of the natural 
history of the coati—subspecific assignment of popu¬ 
lations—that also requires clarification. Historically, 
Bailey (1905) assigned Texas specimens to the sub¬ 
species N. n. yucatanica (Allen 1904; type locality, 
Chichen Itza, Yucatan, Mexico) and that assignment 
remained in effect until Goldman (1942) described 
N. n. tamaulipensis (type locality, Cerro de la Silla, 
near Monterrey, Nuevo Leon, Mexico) and assigned 
the Texas populations to that subspecies. Specimens 


from Arizona and New Mexico were assigned to the 
subspecies N. n. pallida (type locality, Guadalupe y 
Calvo, Chihuahua, Mexico) that also was described by 
Allen (1904). All of these subspecies were combined 
in 1951 by Phillip Hershkovitz into a single subspe¬ 
cies, N. n. molaris (type locality, Manzanillo, Colima, 
Mexico), which had been described in 1902 by Merriam 
and had taxonomic priority. Hershkovitz’s taxonomic 
assignment was based solely on color, reflecting the 
presence of a major shift in color across the Isthmus 
of Tehuantepec. Hofifineister (1986) called for a de¬ 
tailed analysis, employing cranial features as well as 
color and other characters, to clarify the relationships 
of coati populations. We agree with Hoffmeister’s 
assessment but also would suggest a genetic analysis 
should be added to better determine if the Arizona and 
Texas populations are part of the same population or 
represent distinct population clades. 


Acknowuedgments 


We thank the following individuals for providing 
information, photos, or comments about coatis included 
in many of the sightings in our compilation: Javier de 
Leon, Robert Dowler, Chris Durden, Jim Finegan, Bea 
Harrison, Dean Hendrickson, Tony Henehan, Joe Jop¬ 
lin, Jerry Johnson, Rob Klockman, Mark Lockwood, 
David Long, Chris Mallery, Bonnie McKinney, Glenn 
Merkord, Pat Merkord, Jim Mueller, Marcos Paredes, 
Raymond Skiles, Nathan Wells, and Bryan Hughes. 


Thanks go also to Robert Bradley and Jonah Evans 
for reviewing early drafts of the manuscript. George 
Ruffner helped obtain information on the conserva¬ 
tion status of the coati in Arizona. Several individuals 
kindly shared information about camera trap studies, in¬ 
cluded Skyler Stevens, Catherine Dennison, Ray Willis, 
Michael Huston, James Eddy, Marc Cancellare, Robert 
Dowler, and Michael Tewes. Lisa Bradley kindly as¬ 
sisted with the preparation of the map for Figure 2. 


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Addresses of authors: 

David J. Schmiduy 

no. 60 Homesteads Road 
Placitas, New Mexico 87043 
djschmidly @ gmail. com 

John Karges 

The Nature Conservancy 
200 East Grayson Street 
Suite 202 

San Antonio, Texas 78215 
jkarges@tnc.org 


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259:1-15. 


Robert H. Dean 

Department of Interpretation 
Big Bend National Park 
P. O. Box 556 
Terlingua, Texas 79852 
rgvranger@ yahoo, com 


Ecological Distribution and Foraging Activity of the Ghost-faced Bat 
(Mormoops megalophylla ) in Big Bend Ranch State Park, Texas 


Franklin D. Yancey, II 
Abstract 

Data from a two and one-half year study of bats in Big Bend Ranch State Park 
(BBRSP), Texas, along with those from subsequent monitoring work were used to 
ascertain the distribution of the Ghost-faced Bat ( Mormoops megalophylla) within the 
park. In addition, habitat affinities and periods of foraging activities were assessed. 
A GIS-generated map of the distribution of Mormoops megalophylla within BBRSP 
indicated that this species was relatively widespread throughout BBRSP. Mormoops 
megalophylla was found to occur in both flatlands and canyonlands, but favored the 
latter. In addition, the Ghost-faced Bat was found to prefer riparian areas with little or 
no vegetation, as opposed to areas with dense vegetation. Females appear to reside in 
the park only during late spring and summer, whereas adult males apparently are absent 
from the area throughout the year. Prey items of M. megalophylla consisted mostly of 
lepidopterans, with coleopterans, dipterans, hemipterans, homopterans, and neuropter- 
ans being consumed at a much lesser degree. Individuals were found to forage above 
standing or slow-moving water between 2154 h and 0700 h, but were most active the 
first two hours following sunset. 

Key words: Big Bend Ranch State Park, diet, distribution, foraging activity, 
Ghost-faced Bat, habitat, Mormoops megalophylla , Texas 


Introduction 


The Ghost-faced Bat {Mormoops megalophylla) 
ranges from northern South America, northward 
through parts of Central America, on up to northern 
Mexico. It reaches its northern limits in the extreme 
southern United States, where it is known only from 
southern Arizona and southwestern Texas (Beatty 1955; 
Smith 1972; Hall 1981; Hoffmeister 1986; Rezsutek 
and Cameron 1993). Although Mormoops megalo¬ 
phylla is not particularly uncommon throughout much 
of its range, data on the life history of this species are 
nearly unknown. For example, Rezsutek and Cameron 
(1993) summarized the biology of M. megalophylla 


throughout its range, but presented little information on 
its habitat affinities and foraging patterns and activities. 
Yancey (1997) made some general comments regarding 
habitat preferences, but did not quantify foraging habi¬ 
tat affinities. Bateman and Vaughan (1974) presented 
some general information on mormoopid periods of 
activity, however most of their observations were of 
bats traveling to and from foraging areas. This study 
details the distribution of M. megalophylla within Big 
Bend Ranch State Park (BBRSP), Texas, and provides 
insight into the habitat affinities and foraging activities 
of this poorly understood bat. 


147 


148 


Clyde Jones Memorial Volume 


Materials and Methods 


This study was conducted entirely within the 
boundaries of BBRSP, which lies within the Trans- 
Pecos region of the extreme western part of Texas 
(see Fig. 1). The park is located just north of the 
Mexican state of Chihuahua, from which it is sepa¬ 
rated by the Rio Grande. The town of Lajitas and Big 
Bend National Park occur to the east, and the city of 
Presidio is found to the west. The major portion of 
BBRSP is situated in the southeast corner of Presidio 
County, whereas a small part of the park occurs in the 
southwest corner of Brewster County. Initially the 
park consisted of approximately 113,000 ha (Alloway 
1995), but with land acquisitions over the past 20 years 
the park has increased in size to more than 125,000 ha 
(M. W. Lockwood personal communication; unrefer¬ 
enced). Most of the park is composed of scrub habitat 
dominated by typical Chihuahuan Desert plants such 
as Creosote-bush (Larrea tridentata ), acacia ( Acacia 
sp.), Lechuguilla ( Agave lechuguilla ), and a variety 
of cacti ( Echinocereus sp., Mammalaria sp., Opuntia 
sp.). However, with more than 100 springs and several 
permanent streams, the park also contains many locali¬ 
ties with riparian habitat dominated by cottonwoods 
(Populus sp.), willows ( Salix sp.), and seepwillows 
(Baccharis sp.). These numerous water-associated 
habitats within the park were the focal points for sam¬ 
pling bats during this study. 

This study was done concurrently with a general 
assessment of the mammalian fauna of BBRSP that 
occurred from 1994 to 1996 (see Yancey 1997), with 
subsequent follow-up field work conducted in 2015. 
Bats were sampled using mist nets as outlined by Kunz 
and Kurta (1988) and Kunz et al. (1996). At dusk, nets 
were strung across selected springs, streams, stock 
tanks, or other small bodies of water that occur through¬ 
out BBRSP, and monitored throughout the night. Each 
night that an individual net was set and monitored was 
considered one net-night. Localities sampled varied by 
the density of riparian vegetation in the immediate area, 
as well as the surrounding topography. Vegetation was 
considered closed if it formed a closed canopy or was 
dense and tall enough to obstruct the edges of the body 
of water at the net site. If the immediate vegetation was 
short and sparse to the point where the net was unob¬ 


structed and/or no canopy formed, it was considered 
open. Localities were labeled as canyonlands if the site 
was within a canyon greater than 5 m deep, whereas 
they were considered flatlands if the site was situated 
on level terrain or within a shallow arroyo. 

Localities of sample sites were obtained with a 
Magellan hand-held global positioning system (GPS). 
Each individual captured was measured, sexed, and ex¬ 
amined for reproductive condition, and when feasible, 
time of capture was noted. Following collection of data, 
bats were released, or, in some instances, retained as 
voucher specimens; those specimens are deposited in 
the Natural Science Research Laboratory of the Mu¬ 
seum of Texas Tech University. Stomachs (along with 
contents) of some bats were collected and analyzed 
following Whitaker (1988). Data were recorded in an 
Excel-based database and downloaded into Arc View 
GIS software for generation of the distribution map. 

Habitat associations were analyzed by calculating 
chi-square values and then plotting those values against 
a resampled distribution obtained at 1,000 iterations 
(Bruce 1992; Simon 1992). The alpha level was set 
at 0.05. Habitat variables (vegetation density and to¬ 
pography) were analyzed independently and combined. 
Because there were only two treatment groups for each 
independent test, there was no need for further analysis 
if a significant difference was noted. Because there 
were four treatment groups in the combined variable 
test, if a significant difference was detected then a mul¬ 
tiple comparison chi-square test was used to determine 
which habitats differed from one another (Glantz 1992). 
The alpha level of rejection for this test was determined 
using the Bonferroni inequality formula, 

a=a 2 /k 

x 

where a 2 is the original alpha level, and k is the num¬ 
ber of individual treatment groups considered in the 
original test (Glantz 1992). Therefore, for the multi¬ 
comparison test used in this study the alpha level was 
0.0125. A binomial test against equal proportions with 
an alpha level of 0.05 was used to determine peak pe¬ 
riod of activity (Dowdy and Wearden 1991). 


Yancey, II—Ghost-Faced Bat in Big Bend Ranch State Park, Texas 


149 


i_i_i 

KILOMETERS 

Figure 1. GIS-generated map depicting the distribution of the Ghost-faced Bat (Mormoops megalophylla) 
in Big Bend Ranch State Park, Texas. Stars represent localities where M. megalophylla was captured 
and circles indicate localities sampled that were negative for M. megalophylla. The scale of the map is 
such that if all symbols representing localities were plotted, there would be considerable overlap, thus 
confounding the map. Therefore, where overlap would occur, a single symbol has been plotted that 
represents multiple localities in the same general area. 


150 


Clyde Jones Memorial Volume 


Results and Discussion 


Distribution and abundance. —From January 
1994 to June 1996, 339 mist nets were set at 108 lo¬ 
calities throughout BBRSP. A total of 550 bats repre¬ 
senting 14 species was captured during this time, 131 
of which were Ghost-faced Bats. They accounted for 
24% of all bats captured, second in abundance only 
to the American Parastrelle (. Parastrellus hesperus ), 
which accounted for 34% of bats taken. Of the 108 
sites sampled, M. megalophylla was encountered at 16 
sites scattered throughout the park. On 5 September 
2015, a 109 th locality was netted. This site is situated 
in the isolated Solitario region of the park, and was 
not sampled during the original study period due to 
the absence of water sources. During the recent sam¬ 
pling of this new locality, six Ghost-faced Bats were 
captured, bringing the total number of sites in BBRSP 
where M. megalophylla has been documented to 17. 
These results indicate that this bat is relatively com¬ 
mon and widespread in the park. The general pattern 
of distribution of M. megalophylla within BBRSP is 
depicted in Figure 1. 

That Ghost-faced Bats were found to be common 
and widespread in the Big Bend region is in contrast 
to previous reports by Easterla (1973) and Scudday 
(1976), who listed this bat as uncommon in Big Bend 
National Park and BBRSP, respectively. The occur¬ 
rence of M. megalophylla at specific sites is known to 
be highly variable and unpredictable (Ammerman et 
al. 2012), and discrepancies in the previously reported 
abundance of this bat in the area indicate the possible 
existence of temporal fluctuations. On 4, 5, and 6 
September 2015, three sites in BBRSP were sampled to 
gain insight into the stability of this species in the park. 
During this time, six nets were set yielding a total of 
103 bats, 47 of which were Ghost-faced Bats, making 
them the most frequently encountered bats (45.6%) 
during this abbreviated sampling period. The results 
of the follow-up work suggest that the population of 
M. megalophylla at BBRSP has remained stable over 
the past two decades. 

Ghost-faced Bats are reported to reside in Trans- 
Pecos Texas only during the warmer months (Schmidly 
1977; Ammerman et al. 2012), and that was found to be 
the situation at BBRSP during this study. Individuals 


were encountered only between 29 March and 18 Sep¬ 
tember. Interestingly, M. megalophylla resides on the 
Edwards Plateau just to the east during winter (Eads et 
al. 1957; Goetze 1998). This suggests seasonal migra¬ 
tion between these two areas, although this has yet to 
be documented (Ammerman et al. 2012). Of the 184 
individuals examined during this study, 182 were adult 
females, the only two males being juveniles. Most of 
the females were either gravid, lactating, or in post- 
lactating condition (see Yancey 1997). It appears that 
prior to or during migration from wintering grounds, 
females segregate themselves from males, then arrive 
at BBRSP to set up nursery colonies. Young are born 
and nursed during the summer at these nursery colonies, 
and then the onset of fall migration leads to exodus from 
the park in late summer or early autumn. 

Ecological affinities. —Specimens ofM megalo¬ 
phylla were acquired from water-associated sites with 
a variety of surroundings. Because M. megalophylla 
was found to occur in the park only from late March 
through September, only net-nights from this time 
period (n = 260) were considered in the analyses of eco¬ 
logical affinities. Ghost-faced Bats occasionally were 
taken among dense stands of plants that often formed 
a closed canopy, but were more often encountered in 
open areas with sparse vegetation (P < 0.001; Table 

1) . They also were captured in both canyons and level 
flatlands, but were more often netted in the former (P < 
0.001; Table 1). When considering these two features 
collectively (amount of vegetation and topography), a 
difference between observed and expected numbers of 
bats was detected (P < 0.001), and therefore pairwise 
comparisons were made (a = 0.0125 based on the 
Bonferroni inequality adjustment). These comparisons 
indicate that M. megalophylla in BBRSP has affinities 
for the various habitats in the following order (highest 
to lowest): canyons with open/sparse vegetation (Fig. 

2) ; flatland areas with open/sparse vegetation (Fig. 3); 
canyons with closed/dense vegetation (Fig. 4); and 
flatland areas with closed/dense vegetation (Fig. 5; 
Table 2). Each habitat was significantly favored over 
the one below it {P < 0.001). 

Apparently, both density of vegetation and topog¬ 
raphy play important roles in the selection of foraging 


Yancey, II—Ghost-Faced Bat in Big Bend Ranch State Park, Texas 


151 


Table 1. Results for two independent habitat variables; amount of vegetation and topography. Only nets set (n = 
260) and individuals caught (n= 131) from late March through mid-September (1994-1995) were considered, as 
this is the time period that M. megalophylla is known to occur in Big Bend Ranch State Park. 


Vegetation 

Topography 

Sparse 

Dense 

Canyon 

Level 

Net-nights 

111 

149 

91 

169 

Ghost-faced Bats caught 

126 

5 

97 

34 

Ghost-faced Bats caught/net-night 

1.135 

0.034 

1.066 

0.201 


Figure 2. Example of canyonland with open/sparse vegetation. This habitat was the type that 
Mormoops megalophylla had the highest affinity for at Big Bend Ranch State Park, Texas. 


habitat of M. megalophylla in BBRSP, but the former 
seemingly more so than the latter. The habit of avoid¬ 
ing densely vegetated areas probably is in response to 
the high aspect wings of this species, as these relatively 
long wings would be a hindrance in thick vegetation 
(Norberg 1994). This bat’s affinity for canyons prob¬ 
ably is due to the abundance of small caves on the sides 
of many canyon walls at BBRSP. Caves reportedly are 
a primary roosting structure of this bat (Schmidly 1977; 
Graham and Barkley 1984; Ammerman et al. 2012), 
and individuals have been collected from caves in the 
adjacent Mexican states of Coahuila (Baker 1958) and 
Chihuahua (Anderson 1972). Foraging near a night 


roost would reduce the amount of time and energy spent 
traveling from roost to foraging grounds as compared 
to foraging some distance from the roost site. 

It has been suggested that windy conditions may 
reduce the effectiveness of mist nets in capturing bats, 
and thus possibly introduce a sampling bias in favor 
of less windy areas (Kunz and Kurta 1988). The 
canyons sampled during this study certainly provided 
some protection from wind that was not present for the 
flatlands sampled. However, wind does not appear to 
alter the effectiveness of mist nets until wind speed 
reaches 14.5 km per hr (O’Farrell and Bradley 1970). 


152 


Clyde Jones Memorial Volume 


Figure 3. Example of flatland with open/sparse vegetation. This habitat was the second 
most frequently used habitat by Mormoops megalophylla at Big Bend Ranch State Park, 
Texas. 


Figure 4. Example of canyonland with closed/dense 
vegetation. This habitat was used only sparingly by 
Mormoops megalophylla at Big Bend Ranch State 
Park, Texas. 


Yancey, II—Ghost-Faced Bat in Big Bend Ranch State Park, Texas 


153 


Figure 5. Example of flatland with closed/dense vegetation. This habitat was the type that 
Mormoops megalophylla had the lowest affinity for at Big Bend Ranch State Park, Texas. 


Table 2. Results for habitat variables combined. Only nets set ( n = 260) and individuals caught (n = 131) from late 
March through mid-September (1994-1995) were considered, as this is the time period that M. megalophylla is known 
to occur in Big Bend Ranch State Park. 


Vegetation: Topography 


Sparse:Canyon 

Sparse:Level 

Dense:Canyon 

Dense:Level 

Net-nights 

67 

44 

24 

125 

Ghost-faced Bats caught 

93 

33 

4 

1 

Ghost-faced Bats caught/net-night 

1.388 

0.750 

0.167 

0.008 


As nets were not typically set during such conditions 
during this study, a sampling bias due to wind should 
be considered minimal at most. 

Diet and foraging activity. —The stomach 
contents of 45 Ghost-faced Bats were analyzed to de¬ 
termine the food items that M. megolophylla forages 
on in BBRSP. Insects made up 100% of the diet. At 
100% frequency of occurrence, lepidopterans were 
by far the most common type of insect consumed, 
followed by coleopterans (including at least some 
scarabids and some carabids), dipterans, hemipterans, 


homopterans, and neuropterans (Table 3). The majority 
(> 50% by volume) of stomach contents for 44 of the 
45 individuals examined was lepidopteran, whereas 
one individual had coleopterans (carabids and others) 
comprising the majority of its stomach contents. Of the 
45 stomachs examined, 37 contained only lepidopteran 
elements. Black (1974) set a percent frequency of oc¬ 
currence level of 65 or greater to classify a bat species 
as a particular type of insect strategist. With a 100% 
percent frequency of occurrence for lepidopterns, M. 
megalophylla clearly warrants classification as a moth 
strategist at BBRSP. 


154 


Clyde Jones Memorial Volume 


Table 3. Prey items recovered from the stomachs of 45 Ghost-faced Bats {Mormoops megalophylla ) from Big Bend 
Ranch State Park. All prey items recovered were of insect material from six insect orders. 


Insect Order 


Lepidoptera 

Coleoptera 

Diptera 

Hemiptera 

Homoptera 

Neuroptera 

Number of stomachs 
recovered from 

45 

3 

3 

1 

1 

1 

Percent frequency of 
occurrence 

100 

6.7 

6.7 

2.2 

2.2 

2.2 


Bateman and Vaughan (1974) summarized the pe¬ 
riod of activity ofM megalophylla in Sinaloa, Mexico, 
and determined that this bat was active between 1930 
h and 0630 h. However, this assessment was based on 
the capture times of only 20 individuals in nets that, in 
many cases, were left unattended throughout the night 
until 0600-0630 h, in which case time of capture was 
vague at best. In other instances, nets were attended, 
but monitored only until 2230 h at the latest, in which 
case the majority of the night was left unsampled. 
Therefore they were able to establish a broad range 
of activity time, but were unable to comment on peak 
periods of activity. During this study, the capture time 
intervals of 69 individuals taken in 10 nets that were 
monitored throughout the night were recorded. These 


I thank Texas Parks and Wildlife personnel for 
logistical support, including the use of facilities and 
vehicles while doing research at BBRSP. Thanks to 
Mark Lockwood for assistance in sampling bats, David 
Sissom for help in identifying insect fragments, and 
Richard Strauss for statistical advice. This project 
was funded by the Natural Resources Program (David 
Riskind, Director) of Texas Parks and Wildlife De¬ 
partment, and supported, in part, by Reedley College 
(Sandra Caldwell, President). Bats were collected in 


nets were set above standing or slow-moving water 
where bats were visibly noted to be foraging on the 
wing, as is typical of mormoopids (Hill and Smith 
1984). Based on times of capture, M. megalophylla 
was found to be actively foraging from just after dusk 
(2154 h) to just after sunrise (0700 h). The first two 
hours following dusk appear to be the peak period of 
activity for this bat, as it was captured significantly 
more often during this time (56 individuals) than at 
all other times combined (13 individuals; P < 0.001). 
The first two hours following dusk also seems to be 
a peak period of foraging for many other species of 
moth-strategist bats in the southwestern United States 
(Jones 1965; Black 1974). 


BBRSP in accordance with scientific collecting permits 
issued by Texas Parks and Wildlife Department (permit 
numbers SPR-079, SPR-189, 4-94, 25-5, 2015-07). 
Thanks to Robert D. Bradley and Richard W. Manning 
for providing constructive comments on a previous ver¬ 
sion of this manuscript. Finally, I wish to thank Clyde 
Jones for all of his assistance, guidance, inspiration, 
and friendship during the course of research for this 
and related projects. 


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Jones, C. 1965. Ecological distribution and activity periods of 
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devices. Pp. 1-29 in Ecological and Behavioral Methods 
for the Study of Bats (T. H. Kunz, ed.). Smithsonian In¬ 
stitution Press, Washington, D.C. xxii + 533 pp. 

Kunz, T. H., C. R. Tidemann, and G. C. Richards. 1996. Captur¬ 
ing mammals (small volant mammals). Pp. 122-148 in 
Measuring and Monitoring Biological Diversity. Stan¬ 
dard methods for mammals (D. E. Wilson, F. R. Cole, J. D. 
Nichols, R. Rudran, andM. S. Foster, eds.). Smithsonian 
Institution Press, Washington, D.C. xxvii + 409 pp. 

Norberg, U. M 1994. Wing design, flight performance, and 
habitat use in bats. Pp. 205-239 in Ecological Morphol¬ 
ogy (P. C. Wainwright and S. M. Reilly, eds.). University 
of Chicago Press, vii + 367 pp. 

O’Farrell, M. J., and W. G. Bradley. 1970. Activity patterns 
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51:18-26. 

Rezsutek, M., and G. Cameron. 1993. Mormoops megalophylla. 
Mammalian Species 448:1-5. 

Schmidly, D. J. 1977. The mammals of Trans-Pecos Texas 
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Mountains National Park. Texas A&M University Press, 
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Natural History, University of Kansas 56:1-132. 

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Address of author: 

Franklin D. Yancey, II 

Oakhurst Center, Reedley College 
40241 Hwy 41 
Oakhurst, CA 93644 
frank.yancey @ scccd. edu 


Pleistocene/Holocene Faunas from the Trans-Pecos 


Arthur H. Harris 


Abstract 

There are only two extensive Pleistocene fossil faunas known from Trans-Pecos 
Texas: Fowlkes Cave and Sierra Diablo Cave. The fauna from Sierra Diablo Cave is 
compared to those from Fowlkes Cave and the Guadalupe Mountains. Both the Fowl¬ 
kes Cave and Sierra Diablo Cave faunas are shown to be mixed Holocene-Pleistocene 
deposits. 

Key words: faunas, fossil, Holocene, Pleistocene, vertebrates, Wisconsin age 


Introduction 


The Guadalupe Mountains represent the south¬ 
ernmost high ground more or less in continuity with 
the southern Rockies of northern New Mexico. Ranges 
immediately to the south are of notably lower eleva¬ 
tion. Wisconsin-age faunas are well documented from 
the Guadalupe Mountains region of southeastern New 
Mexico and adjacent Trans-Pecos Texas. Until recently, 
only Fowlkes Cave (Dalquest and Stangl 1984) had 
produced extensive faunal remains of this age from 
south of that region. Sierra Diablo Cave, roughly 75 
km W of Fowlkes Cave, recently produced a large 
fauna. The purpose of this paper is to document the 
Sierra Diablo Cave fauna and its contribution to our 
knowledge of late Pleistocene paleobiology in relation 
to the Guadalupe Mountains late Pleistocene sites and 
to the Fowlkes Cave fauna (Fig. 1). 

The Late Pleistocene in southeastern New Mexico 
and adjacent Trans-Pecos Texas. —The Wisconsin is 
the last glacial age, lasting from about 75 kya (thou¬ 
sands of years ago) until 11.7 kya. The Wisconsin has 
been divided into early, middle, and late phases. The 
mid Wisconsin is considered to have ended at about 
29,000 kya. 

A variety of late Pleistocene sites occur in the 
Guadalupe Mountains region of southeastern New 


Mexico and adjacent Texas. The most extensive of 
these, Dry Cave in Eddy County, New Mexico, is used 
herein for comparison with sites to the south (Morgan 
and Harris 2015). Radiocarbon dates from the younger 
Dry Cave sites range from 10,730 ± 150 to 15,030 ± 
210 BP (Harris 1989), with older but undated deposits 
extending into the middle Wisconsin. 

Sierra Diablo Cave. —Sierra Diablo Cave is a 
small horizontal cave in the limestone rim rock of the 
Diablo Plateau (Fig. 2). Elevation is approximately 
1,645 m and the exposure is to the south. The Rio 
Grande is about 50 km to the southwest. Seventeen 
Draw, which may have had permanent water during the 
Pleistocene, approaches to within about 1.5 km of the 
cave. The landscape supports typical desert vegetation 
of the region. 

Although snakes and some small mammals 
almost certainly voluntarily utilized the cave, most 
specimens likely were brought in by predators and 
scavengers. The rugged topography would make it 
difficult for horses and the larger artiodactyls to access 
the cave; however, presence of at least two large felids 
is sufficient to explain their presence. Smaller predators 
and avian scavengers undoubtedly account for many 
of the other remains. 


157 


158 


Clyde Jones Memorial Volume 


Figure 1. Location of fossil sites in relation to several political boundaries and place 
names. 


Figure 2. Sierra Diablo Cave is located a bit to the right of center, near the base of the massive limestone 
outcrop. The foreground vegetation is largely creosotebush. View approximately north. Photograph by 
A. H. Harris. 


Harris—Pleistocene/Holocene Faunas from the Trans-Pecos 


159 


Materials and Methods 


A small collection of faunal remains was donated 
to the University of Texas at El Paso (UTEP) from 
presumably surface collections in 1966. More recently, 
the owners of Circle Ranch invited me to initiate 
faunal studies at the cave, and exploratory collections 
were hand-picked or sieved from disturbed matrix in 
2007 and 2009. More formal collections were made 
under a memorandum of understanding between the 
ranch owners and UTEP. Because of the presence of 
archaeological material, formal excavation has been by 
archaeologists. University of Texas at El Paso archae¬ 
ologist David Carmichael held a field school at the site 
in 2010, and Javier Vasquez continued with field parties 
in 2011-2013; all faunal material during these years 
was recovered by the archaeologists during screening 
for artifacts. Small matrix samples also were taken. 
Most faunal material from 2011-2013 was collected by 
hand by the archaeological field crews from %-inch or 
window-screen mesh. 

To date, I have identified and catalogued 1,666 
specimens into the UTEP Biodiversity Paleobiology 
Collection. These form the basis for interpretation of 
the Sierra Diablo Cave fauna. 

Stratigraphy and chronology .—Basic stratigraph¬ 
ic information was established by Vasquez (2010). The 
cave has been extensively pot hunted, with excavations 
extending into Pleistocene deposits. Much of the fauna 
comes from an overburden of disturbed sediments. 
Although various sizes of limestone roof-fall occur, 
the basic matrix is of powdery fill, possibly of eolian 
origin. Because of pot hunting, aboriginal activity, 
the softness of the fill, and animal burrowing, mixing 
between levels has occurred, especially in the upper 
portions. During excavation, attempts were made to 
separate disturbed overburden from the original sur¬ 
face, but the division point was not always apparent. 
Faunal material labeled by the archaeologists as from 
disturbed areas is given separately in Table 1; material 
not so labeled is given as reported by the excavators, 
but may not have been recognized as from disturbed 
sediments by the archaeological crew. 

A datum level was established 0.69 m above 
ground surface. The archaeological excavations have 


revealed seven recognizable strata, or zones: A through 
G (uppermost to lowest). Zones E and G are limited in 
scope; the latter lacked fossil remains and is not further 
considered here. Several of the strata were subdivided 
into levels by the excavators. 

The surface fill (A), excavated in three levels and 
ranging from 0.54 to 0.75 m below datum (thicknesses 
measured at the same point for all strata), contained 
much plant material and produced artifacts of archaic 
cultural age as well as of the 20 th century; however, 
occurrences of Pleistocene taxa such as horse (Equus) 
and Conkling’s Pronghorn (Stockoceros conklingi ) 
indicate strong contamination (Table 1). Judging from 
the fauna, zone A (Holocene, based on cultural material 
and presence of modern fauna) may lie on a Pleistocene 
surface (zone B) with much mixing between A and the 
upper portions of B. 

Stratum B ranged from 0.76 to 1.03 m below da¬ 
tum. Consisting largely of compacted sediments, there 
was little in the way of plant material. Some cultural 
material was present. Stratum C ranged in depth from 
1.03 to 1.54 m below datum. There is a sharp break 
between B and C, the latter being a loose deposit of 
coarse silt to fine-grained sand. Some cultural material 
was present. 

Stratum D, 1.54 to 1.88 m below datum, consisted 
of silt-sized particles. The only cultural material pres¬ 
ent was flake debitage. Stratum E was shallow and 
pinched out in places, allowing stratum D to contact 
stratum F. Except for color, composition was essen¬ 
tially the same as F, which consisted of carbon-stained 
sediments ranging from 1.93 to 2.06 m below datum. 

A radiocarbon date of 32,770 ±38 years BP 
(AA94457) was determined on charcoal from stratum 
F. Because of excavation conditions at the time, it is 
unclear whether the date actually appertains to stratum 
F, but supports the presence of mid Wisconsin deposits. 
The only other date available to me is 34,970 ± 640 
(about 40,000 radiocarbon kya) years BP (AA97732), 
determined on charcoal from stratum F. The sample 
was taken 30 cm from a bone of an extinct pronghorn 
and 20 cm from a chert biface. Under microscopic 


160 


Clyde Jones Memorial Volume 


Table 1. List of taxa recovered from the late Pleistocene of the Guadalupe Mountains region (GM), the older fossil 
deposit of Fowlkes Cave, and the Holocene to mid Wisconsin of Sierra Diablo Cave by stratigraphic zone. “Disturb” 
indicates taxa from disturbed sediments of Sierra Diablo Cave; “cf.” indicates a degree of uncertainty in identifica¬ 
tion; “?” indicates a questionable identification; indicates presence in the terminal Pleistocene only; | indicates 
an extinct species. Birds, unrecorded from Fowlkes Cave, are placed last in the table and only those present in Sierra 
Diablo Cave are compared to those from the GM. Nomenclature of Dalquest and Stangl (1984), Parmley (1988,1990), 
and Parmley and Bahn (2012) is updated to current usage (original names in parentheses). 


Taxon 

GM 

Fowlkes 

A 

B 

c 

D 

E 

F 

Disturb 

Osteichthyes 

X 


X 

Arnbystoma mavortium 

X 


X 


X 

Anaxyrus 

X 


X 

X 


X 

X 

X 

Anaxyrus cognatus 

X 


Anaxyrus punctatus 

X 


X 

X 

Anaxyrus woodhousii (Bufo woodhousei ) 

X 

X 


Anaxyrus microscaphus/woodhousii 

X 


cf. 


Incilius nebulifer (Bufo valliceps) 


X 


Pseudacris triseriata 

X 


Lithobates ( Rana ) 

X 

X 


? 

Lithobates pipiens 

X 


Scaphiopus/Spea 

X 


X 


Scaphiopus couchii 

X 


cf. 


X 

Spea bombifrons 

X 


cf. 

Spea multiplicata 

X 


X 

Terrapene 

X 


cf. 


Gopherus morafkai 


X 

X 


X 


X 

X 

Crotaphytus collaris ( Crotaphytus ) 

X 

X 

X 

X 

X 

X 

X 

X 

X 

Gambelia wislizenii 


? 

Holbrookia maculata 

X 


Phrynosoma cornutum 

X* 

X 


X 


X 

X 

Phrynosoma hernandesi 

X 


X 

X 

X 


X 


X 

Phrynosoma modestum 

X* 

X 

X 

X 


X 

X 


X 

Sceloporus 

X 


X 

cf. 

cf. 


X 

X 


Sceloporus cowlesi x 

Plestiodon cf. P. obsoletus {Eumeces cf. E. 
obsoletus) 


Plestiodon multivirgatus 


x 


Harris—Pleistocene/Holocene Faunas from the Trans-Pecos 


161 


Table 1. (cont.) 


Taxon 

GM 

Fowlkes 

A 

B 

c 

D 

E 

F 

Disturb 

Aspidoscelis ( Cnemidophorus ) 

X* 

X 

X 


Arizona elegans 


X 

X 


X 

Bogertophis subocularis 

X 

X 

X 

X 


X 


X 

X 

Coluber/Masticophis 

X 

X 

X 

X 

X 


X 

X 

Gyalopion canum 


X 


Lampropeltis alterna {L. mexicana ) 


X 


Lampropeltis getula 

X 

X 

X 

X 

X 


X 

Lampropeltis triangulum 


X 


Opheodrys aestivus 


X 


Pantherophis emoryi (Elaphe cf. E. guttata ) 

X 

X 

X 

X 


X 

Pituophis catenifer (P. melanoleucus ) 

X 

X 

X 

X 

X 

X 


X 

Rhinocheilus lecontei 


X 

X 

X 


cf. 


X 

Salvadora 

X 


X 

X 


X 

Sonora 


X 


Tantilla 


X 


Trimorphodon vilkinsonii 


X 

Crotalus 

X 

X 

X 

X 

X 


X 

X 

Crotalus atrox 

X 


cf. 


Diadophis punctatus 


cf. 


Heterodon nasicus 


X 

Hypsiglena 

X 


Hypsiglena jani (H. torquata ) 


X 


Nerodia erythrogaster 


X 


Thamnophis 

X 

X 


Thamnophis marcianus 


cf. 


t Nothrotheriops shastensis 

X 


X 


Cynomys 

X 

X 

X 

X 

X 


X 

X 

X 

Cynomys gunnisoni 

X 


X 


cf. 

Cynomys ludovicianus 

X 

X 

X 

X 


X 

Ictidomys/Xerospermophilus 

X 

X 

X 

X 


X 

X 

Ictidomys tridecemlineatus 

X 


Xerospermophilus spilosoma (Spermophilus 
spilosoma) 


X 


162 


Clyde Jones Memorial Volume 


Table 1. (cont.) 

Taxon 

Marmota flaviventris 

Otospermophilus variegatus {Spermophilus varie- 
gatus ) 

Tamias ( Eutamias ) 

Urocitellus elegans 
Chaetodipus 
Chaetodipus hispidus 
Dipodomys merriami/ordii 
Dipodomys merriami 
Dipodomys ordii 
Dipodomys spectabilis 
Perognathus 

Perognathus flavus/merriami 
Perognathus flavus 

Cratogeomys castanops {Pappogeomys 
castanops ) 

Geomys arenarius 

Thomomys bottae 

Thomomys talpoides 

Lemmiscus curtatus 

Microtus 

Microtus longicaudus 

Microtus mogollonensis {Microtus mexicanus ) 

Microtus ochrogaster 

Microtus pennsylvanicus 

Ondatra zibethicus 

Neotoma cinerea 

Neotoma floridana 

Neotoma leucodon {Neotoma albigula ) 

Neotoma mexicana 
Neotoma micropus 

Onychomys arenicola {Onychomys torridus ) 
Onychomys leucogaster 


GM 

Fowlkes 

A 

B 

c 

D 

X 

X 

X 

X 


cf. 

X 


X 


X 


X 


X 


X 

X 


X 

cf. 


X 

X 


X 

X 


X 


X 


X 

X 


X 

X 

X 


X 

X 

X 

X 

X 


X 

X 

cf. 


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 


F Disturb 

x 

X X 


X X 


X 


X X 


X X 


X X 

X 


X X 


X X 


X 


X 


X X 


cf. 

X 


Harris—Pleistocene/Holocene Faunas from the Trans-Pecos 


163 


Table 1. (cont.) 

Taxon 
Peromyscus 
Peromyscus boylii 
Peromyscus crinitus 
Peromyscus eremicus 
Peromyscus leucopus 
Peromyscus maniculatus 
Peromyscus nasutus (P. difficilis ) 
Peromyscus laceianus (P. pectoralis) 
Reithrodontomys 
Reithrodontomys fulvescens 
Reithrodontomys megalotis 
Reithrodontomys montanus 
Sigmodon hispidus 
Sigmodon ochrognathus 
Erethizon dorsatum 
t Aztlanolagus agilis 
Lepus 

Lepus californicus 
Lepus townsendii 
Sylvilagus audubonii 
Sylvilagus fioridanus/robustus 
Sylvilagus nuttallii 
Cryptotis parva 
Notiosorex crawfordi 
f Notiosorex dalquesti 
t Notiosorex harrisi 
Sorex merriami 

Sorex neomexicanus ( S. vagrans) 
Sorex palustris 
Sorex preblei 
t Desmodus stocki 
Tadarida brasiliensis 


GM Fowlkes A 


C D 


F Disturb 


x 

cf. 

x 

X 

X 

X 


X 

X 

X 

X 

cf. 

X 

X 


X 

X 

X 

X 

X 

X 

X 

X 

cf. 

X 

X 

X 

X 

X 

X 

X 

X 

X 


X 

X 


X 

X 


X 

X 


X 

cf. 

cf. 


X 

X 


X X 


X 

X 


cf. 


X 

X 

X 

X 

X 

X 

X 

X 

cf. 


cf. 


X 

X 


164 


Clyde Jones Memorial Volume 


Table 1. (cont.) 

Taxon 

GM 

Fowlkes 

A 

B 

c 

D 

E 

F 

Disturb 

Antrozous pallidus 

X 


X 

X 


cf. 


cf. 

X 

Corynorhinus townsendii 

X 


X 


Eptesicus fuscus 

X 

X 

X 


X 


X 

Lasiurus cinereus 

cf. 


Myotis californicus 

X 


Myotis californicus/ciliolabrum 

X 


Myotis evotis 

X 


Myotis lucifugus 

X 

X 


Myotis thysanodes 


X 

X 


Myotis velifer 

X 

X 


Lynx rufus (Felis rufus) 

X 

X 

X 

X 

X 

X 


X 

X 

^Panthera atrox 

X 


X 

Puma concolor 

X 


X 

X 


X 

X 

Canis latrans 

X 


X 

X 


X 

X 

Canis dirus/lupus 


X 

Canis dims 

cf. 


Canis lupus 

X 


Urocyon cinerioargenteus 

cf. 


cf. 

cf. 


Vulpes macrotis/velox 

X 


X 

cf. 


X 

Vulpes macrotis 

X 


Vulpes velox 

X 


Ursus americanus 

X 


X 

Mustela erminea 

X 


Mustela frenata 

X 


X 

Taxidea taxus 

X 


X 


X 

X 

Mephitis 

X 


cf. 

X 

cf. 

X 


X 

X 

Spilogale gracilis/putorius 

X 


X 


X 

X 

Bassariscus astutus 

X 


cf. 


X 

Equus 

X 


X 

X 

X 

X 


X 

X 

^ Equus conversidens 

X 


X 


X 


t Equus francisci x 

t Equus occidental'is 


x 


Harris—Pleistocene/Holocene Faunas from the Trans-Pecos 


165 


Table 1. (cont.) 


Taxon 

GM 

Fowlkes 

A 

B 

c 

D 

E 

F 

Disturb 

t Equus scotti 

X 


cf. 

t Mylohyus 


X 


Came lops hestemus 

X 


t Hemiauchenia macrocephala 

X 


X 

Odocoileus 

X 


X 

cf. 


X 

Odocoileus hemionus 

cf. 


X 


X 

Antilocapra americana 

X 


X 

X 


X 

X 

t Capromeryx furcifer 

X 

X 

X 

X 


X 


X 

X 

t Stockoceros conklingi 

X 


X 

X 

X 

X 


X 

X 

t Bison antiquus 

cf. 


cf. 


cf. 

t Oreamnos harringtoni 

X 


X 

Ovis canadensis 

X 


X 


X 

cf. 

Callipepla 

cf. 


X 

t Meleagris crassipes 

X 


cf. 


Cathartes aura 


cf. 


? 

X 

t Coragyps occidentalis 

X 


X 

X 

X 

X 

cf. 


X 

t Gymnogyps ampins 

X 


cf. 

X 


cf. 

X 

Geococcyx californianus 

X 


X 


Bubo virginianus 

X 


X 

Falco sparverius 

X 


X 


X 

X 

Corvus corax 

X 


X 

X 


X 

Corvus cryptoleucus 


X 


Pica hudsonia 

X 


X 

Petrochelidon fulva/pyrrhonota 


X 

X 


X 


X 

X 


examination, the sample was found to be composed 
of plant remains and rodent dung. This suggests that 
the material was a packrat midden used as fuel. Thus, 
although the age of the midden was -35,000 14 c years 
old, the date of the burning could have been much later. 

Occurrence of some cultural material, though 
decreasing with depth, suggests some contamination 
through at least stratum D, and probably below. 


Pleistocene faunal elements are widespread with¬ 
in the sediments, and discovery of the extinct Stock’s 
Vampire Bat (Desmodus stocki ) and the extinct Aztlan 
Rabbit (Aztlanolagus agilis ), species believed to have 
become eradicated from the Southwest by the end of 
the mid-Wisconsin, indicate deposition commenced 
before the late Wisconsin. 

Fowlkes Cave .—Fowlkes Cave is critical for 
interpretation of the Pleistocene faunas south of the 


166 


Clyde Jones Memorial Volume 


Guadalupe Mountains in the western Trans-Pecos. 
The Pleistocene fauna of Sierra Diablo Cave differs 
significantly from that of Fowlkes Cave (Table 1). 
Dalquest and Stangl (1984) interpreted the recovered 
fauna as indicating environmental conditions such that 


microhabitats suitable for taxa now found only far to 
the north were contemporaneous with warm desert 
conditions. This interpretation is incompatible with the 
interpretation of the Sierra Diablo Cave fauna. 


Selected Species Accounts for Sierra Diablo Cave 


Order of presentation follows the Center for 
North American Herpetology for herptiles, American 
Ornithologists’ Union Birds of North and Middle 
America (http://checklist.aou.org/) for birds, and Wil¬ 
son and Reeder (2005) for mammals. f indicates an 
extinct species. 

Crotaphytus collaris (Eastern Collared Liz¬ 
ard).—This is the most common lizard recovered. It 
was present in all zones. Parmley and Bahn (2012), 
with an abundance of caution, took Crotaphytus only 
to the generic level at Fowlkes Cave; I have assumed 
that the taxon represented is of this species and record 
it as such in Table 1. 

The species is widespread in the Southwest today 
and is common in southwestern mid and late Wiscon¬ 
sin fossil faunas, though not recorded in the higher- 
elevation faunas of the Guadalupes. Both rocks and 
some open areas appear to be requirements, according 
to Applegarth (1979). 

Phrynosoma cornutum (Texas Horned Liz¬ 
ard).—This is a warm-climate lizard absent from the 
late Wisconsin Dry Cave faunas until warming at the 
very end of the Pleistocene. It also occurs in the mid 
Wisconsin fauna. It was recovered from zones B and F. 

Phrynosoma hernandesi (Mountain Short-homed 
Lizard).—This homed lizard is by far the most common 
of the genus, with 44 identified elements compared 
to three Texas Horned Lizards and 13 Round-tailed 
Horned Lizards. Occurrences were in zones A, B, C, 
and E. 

It is absent from the desert-scrub lowlands, but 
occurs in higher grasslands into open forest. It is dis¬ 
tributed from southern Canada south well into Mexico. 


Applegarth (1979) hypothesized that only this 
species of lizard was able to thrive at Dry Cave un¬ 
der full glacial conditions because it is ovoviparous, 
retaining the eggs internally and giving birth to living 
young. Warming of body and eggs is accomplished 
by behavioral means (e.g., basking), whereas soil 
temperatures in much of their range may be marginal 
for egg development. 

Occurrence appears continuous through mid and 
late Wisconsin faunas in southern New Mexico, thus 
suggesting that summer temperatures were relatively 
cool throughout the mid and later Wisconsin. 

Phrynosoma modestum (Round-tailed Horned 
Lizard).—This is another warmclimate horned lizard 
apparently absent from all but the Guadalupe Moun¬ 
tains terminal Pleistocene deposits. It was recovered 
from zones A, B, D, and E. 

Aspidoscelis sp. (whiptail lizards).—Although 
present rarely in the mid Wisconsin of the Guadalupe 
Mountain sites, whiptail lizards apparently were extir¬ 
pated during the late full glacial, reentering the region 
only at the tail end of the Wisconsin. It was identified 
from zone A. 

Gopherus morafkai (Morafka’s Desert Tor¬ 
toise).—This tortoise currently is limited to the Sonoran 
Desert; however, it is known from several localities and 
times within the Chihuahuan Desert (Van Devender et 
al. 1976; Harris 2003). At least two eastward pulses 
occurred, one in mid Wisconsin time and one in late 
Wisconsin. Remains were recovered from zones A, B, 
D, and F. The stratigraphic occurrences at Sierra Diablo 
Cave indicate that the mid Wisconsin is represented; 
occurrences in B suggest possible presence in the late 
Wisconsin, also. 


Harris—Pleistocene/Holocene Faunas from the Trans-Pecos 


167 


This tortoise appears to be limited geographically 
today by harsh winter temperatures, implying winters 
at the times of occurrence lacked the extreme cold 
outbreaks seen today. Contemporaneous vegetation 
from Late Wisconsin woodrat middens at Shelter Cave 
in south-central New Mexico indicates the tortoise there 
was living in xerophilous woodlands (Van Devender 
etal. 1976). 

f Nothrotheriops shastensis (Shasta Ground 
Sloth).—This rather regionally common sloth is repre¬ 
sented in the fauna by a single tooth fragment from zone 
A. These sloths seemingly utilized caves for shelter 
and thus remains commonly are found in caves that 
have accessible entrances. They would be expectable 
in caves with vertical entrances only by rare accident. 

A study by McDonald and Jefferson (2008:321) 
suggested that the “lower limiting temperature falls in 
the range of 10 to 20°C,” thus indicating mild winter 
temperatures. It appears that these sloths were absent 
in the Guadalupe Mountains region during full glacial 
times, but present in the mid Wisconsin and again after 
full-glacial conditions. 

Cynomys gunnisoni (Gunnison’s Prairie Dog).— 
The cave currently is within the ranges of C. ludovi- 
cianus, the Black-tailed Prairie Dog, with a presence 
only a few hundred meters away. The only specimen 
in presumed undisturbed sediments was retrieved from 
zone B. Gunnison’s Prairie Dog is known as a late 
Wisconsin fossil at sites in the region. The nearest 
contemporary occurrence is in the northwestern third 
of New Mexico, where it may thrive at relatively high 
elevations. 

Cynomys ludovicianus (Black-tailed Prairie 
Dog).—Occurrence was limited to zones A and B. 
Several specimens were from the surface. In general, 
Black-tailed Prairie Dogs require greater expanses of 
open, low vegetation than does Gunnison’s Prairie Dog. 

Tamias sp. (chipmunk).—Chipmunks have not 
been identified from the extensive Dry Cave late Wis¬ 
consin faunas, while present in the earlier Wisconsin 
fauna and the mid Wisconsin Big Manhole Cave (Mor¬ 
gan and Harris 2015). They have been identified from 
the late Wisconsin of two higher elevation caves (ca. 


2000 m) in the Guadalupe Mountains (Logan 1983; 
Harris and Hearst 2012). Seemingly, they occurred 
only in the higher elevations during the late Wisconsin, 
but descended to lower sites in the mid Wisconsin. 
However, the gray-footed chipmunk currently does 
occur in the Sierra Diablo and Guadalupe Mountains 
(Schmidly 1977). 

Marmota flaviventris (Yellow-bellied Mar¬ 
mot).—Now occurring no closer than northern New 
Mexico, this large sciurid was wide-spread over the 
Southwest in the late Pleistocene. Lundelius (1979) 
suggested that this taxon (and the Bushy-tailed Wood- 
rat, Neotoma cinereus ) lingered on into the Holocene 
in a mesic canyon of the Guadalupe Mountains. It 
also is possible that the remains are Pleistocene in age. 

Remains were recovered from zones A and B. It 
is hypothesized that current absence from sites south 
of northern New Mexico is due scanty winter precipita¬ 
tion and the long spring drought common to the region, 
resulting in lack of green fodder during the time the 
marmots awaken from hibernation (Harris 1970). 

Dipodomys merriami (Merriam’s Kangaroo 
Rat).—Dalquest and Stangl (1984) reported 14 D. 
merriami jaws from Fowlkes Cave. As they noted, 
“Wherever found, D. merriami is an indicator of true, 
arid, desert conditions” (p. 443). Although a number 
of identifications of D. merriami/ordii (that is, either 
one or the other species) have been made within the 
region, there is not one other identification specific 
to this species from Pleistocene levels. Both the two 
specifically identifiable specimens from Sierra Diablo 
Cave are from zone A and obviously Holocene. Thus 
it appears that D. merriami is a true indicator of the 
Holocene in our region. 

Cratogeomys castanops (Yellow-faced Pocket 
Gopher).—Whenever three species of pocket gophers 
occur in a region (as apparently was the case at Sierra 
Diablo Cave), the landscape is divided into mutually 
exclusive tracts. This species tends to inhabit relatively 
deep, silty or sandy soils with few rocks, whereas 
Geomys takes to sandy soils and Thomomys to shallow 
rocky soil. In the absence of other species, the deeper 
soils may be inhabited by any species. 


168 


Clyde Jones Memorial Volume 


This species seems well adapted to relatively arid 
habitats and is widespread in mid and late Wisconsin 
faunas of the region. Its presence implies relatively 
deep soils within predator range, perhaps along Sev¬ 
enteen Draw. At Sierra Diablo Cave, it was recovered 
from all zones except E. 

Geomys cf. arenarius (Desert Pocket Gopher).— 
The specific designation is on the basis of current dis¬ 
tribution; G. knoxjonesi (Jone’s Pocket Gopher) ranges 
fairly close and is an alternative possibility. No Geomys 
occurrences are reported from the eastern side of the 
Guadalupe Mountains region, but the genus is common 
at Pendejo Cave to the west. At Sierra Diablo Cave, 
it is recorded from all zones. Occurrence on the flats 
immediately south of the cave is suggested. 

Thomomys bottae (Botta’s Pocket Gopher).— 
Thomomys bottae was rare. Only one specimen is 
relatable to a zone (zone B), although seven specimens 
are known from disturbed areas. This is a common 
species in a number of late Wisconsin regional sites and 
common in the area today. Current conditions suggest 
that soils likely were relatively shallow and rocky on 
the flats above the cave and inhabitable by T. bottae. 

Thomomys talpoides (Northern Pocket Go¬ 
pher).—Only a single specimen identifiable as T. 
talpoides (plus a queried identification) has been re¬ 
covered. The sample containing the specimen spanned 
two zones: C and D. The species is not known from 
the mid Wisconsin of the Guadalupe Mountains area. 

Since two species of pocket gopher are almost 
always allopatric on a fine scale, presumably different 
ecological habits were inhabited by the two species of 
Thomomys. One likelihood is that the cooler northern 
slopes were preferred by T. talpoides ; since this habitat 
is lacking near the cave, the scarcity of T. talpoides is 
explained. 

Lemmiscus curtatus (Sagebrush Vole).—Al¬ 
though the number of microtine rodents from the site 
is small, the absence of this vole could be significant. 
This is one of the more common voles in the late Wis¬ 
consin Dry Cave sites, making up 37% of the voles 
identified from the Balcony Room site, for example. 
Although not strictly limited to sagebrush grasslands, 


the vole is commonly associated with sagebrush; its 
absence suggests Sierra Diablo Cave may be south of 
late Wisconsin sagebrush distribution. 

Microtus mogollonensis (Mogollon Vole).—This 
is the most forgiving of dry conditions of the regional 
voles, in places descending down into pinyon-juniper 
habitat. If Sierra Diablo Cave was of a somewhat 
more arid aspect than the Guadalupe region to the 
north, it would make sense that Microtus longicaudus 
(Long-tailed Vole) would be absent. Ten of 11 identifi¬ 
able Microtus elements recovered from Sierra Diablo 
Cave are of this species. This vole survived well into 
the Holocene in the Davis Mountains (Kennedy and 
Jones 2006). 

Microtus pennsylvanicus (Meadow Vole).—A 
single partial palate with left Ml and M2 from over¬ 
burden sediments was identified on the basis of a fifth 
“button-shaped” element at the posterior end of M2 
(Semken and Wallace 2002). Also, a visible suture 
between the maxillary and palatine rules out M. mogol¬ 
lonensis (which, alone among southwestern species of 
Microtus , has these elements fused). The assumption 
is made that species, such as M. longicaudus, that 
require more mesic conditions and occasionally show 
the “button,” are absent; this is strengthened by only M. 
mogollonensis being represented among other identifi¬ 
able elements. 

Although the Meadow Vole occurs in mesic mon¬ 
tane habitats, it also inhabits lower-elevation marshy 
areas. Occurrence at Sierra Diablo Cave may indicate 
former habitat in Seventeen Draw or, failing that, in the 
Rio Grande Valley to the southwest. The latter retained 
a population in southern New Mexico at least into the 
early Holocene (Smartt 1977), and there is no reason to 
not suspect occurrence in the Rio Grande Valley nearer 
to Sierra Diablo Cave. 

Neotoma cinerea (Bushy-tailed Woodrat).—This 
northern woodrat is nearly ubiquitous in late Wisconsin 
sites in the region. It was present in zones A, B, C, and 
D. It may have survived into the Holocene in a mesic 
canyon in the Guadalupe Mountains (Lundelius 1979), 
but there is no indication of survival into the Holocene 
elsewhere in the Southwest. 


Harris—Pleistocene/Holocene Faunas from the Trans-Pecos 


169 


Neotoma mexicana (Mexican Woodrat).—These 
woodrats occurred in prepleniglacial and post-plenigla- 
cial deposits at Dry Cave, but apparently were absent 
during the full glacial impact. They also survived into 
the early Holocene at Dry Cave. It was represented 
by a single element at Fowlkes Cave (Dalquest and 
Stangl 1984), but has not been identified from Sierra 
Diablo Cave. 

Onychomys arenicola (Mearn’s Grasshopper 
Mouse).—This mouse is found in a warm-climate, arid 
habitat. It appears in the zones A and B at Sierra Diablo 
Cave and is the current common species of the area. 
It does not appear in the Dry Cave faunas, but does at 
Fowlkes Cave (as Onychomys torridus. Southern or 
Long-tailed Grasshopper Mouse). 

Peromyscus crinitus (Canyon Mouse).—Canyon 
mice occur in the arid West east to northwestern New 
Mexico at present. It has been identified as a fossil in 
the Guadalupe Mountains region (Morgan and Harris 
2015). A single specimen is tentatively identified from 
disturbed sediments at Sierra Diablo Cave, but might 
represent the deer mouse ( Peromyscus maniculatus). 

Peromyscus eremicus (Cactus Mouse).—This is 
a lowland mouse unidentified for sure from the Gua¬ 
dalupe region late Wisconsin sites. An identification 
from Upper Sloth Cave in the Guadalupe Mountains 
apparently was based solely on the absence of acces¬ 
sory cusps (Logan and Black 1979), a trait known to 
occur in other species of Peromyscus. A single Sierra 
Diablo Cave specimen was identified by discriminant 
analysis from zone A. 

Reithrodontomys megalotis (Western Harvest 
Mouse).—Unfortunately, only two specimens of Re¬ 
ithrodontomys of a total of five are identified to species. 
The western harvest mouse is widespread geographi¬ 
cally and ecologically. 

Sigmodon hispidus (Hispid Cotton Rat).—This 
is a relatively warm-climate rodent absent from the 
Guadalupe Mountains late Wisconsin sites until close 
to the terminal Pleistocene. However, west of the Gua¬ 
dalupes, Pendejo and U-bar caves produced both mid 
and late Wisconsin records. This rodent should occur 


near Sierra Diablo Cave at present and is recorded from 
zones A, B, E, and F. 

t Aztlanolagus agilis (Aztlan Rabbit).—This 
small leporid apparently became extinct before the 
height of late Wisconsin full-glacial conditions (Russell 
and Harris 1986), though specimens from U-bar and 
Dust caves could possibly be late Wisconsin. At Dry 
Cave, this rabbit is associated with radiocarbon dates 
between about 25 and 33 kya, though those dates are 
on bone carbonate and likely too young. At Pendejo 
Cave, it disappears after zone K, which likely places 
disappearance at a bit younger than 41 kya. Specimens 
seem fairly securely associated with level 2 of zone B 
and level F. 

The chronological range of this species spans the 
Pleistocene until its extinction. It ranged geographi¬ 
cally from eastern Arizona to central Texas and from 
Colorado into Chihuahua. 

Sorex palustris (Water Shrew).—Despite large 
samples of shrews regionally, the water shrew is re¬ 
corded only from Muskox and Fowlkes caves in the 
region. As the name indicates, this shrew is associated 
with water and high elevation streams in the Southwest. 
Nearest present-day approaches are the high mountains 
of eastern Arizona and northern New Mexico. 

f Desmodus stocki (Stock’s Vampire Bat).—Very 
similar to the living D. rotundus except for its larger 
size, D. stocki survived well into the Holocene on 
San Miguel Island off the coast of California (Guthrie 
1998). However, mainland southwestern records ap¬ 
parently are all mid Wisconsin or earlier. Two of the 
four specimens (all partial humeri) from Sierra Diablo 
Cave are assignable to stratigraphic levels: B, level 4, 
and “B or possibly C?”. This would seem to place level 
B1 as mid Wisconsin. This bat has also been recorded 
in the Trans-Pecos at Terlingua (Cockerell 1930) and 
in New Mexico at U-Bar Cave (Harris 1987). 

Judging from the temperature tolerances of the 
living Desmodus rotundus (Common Vampire Bat), 
presence indicates relatively mild climatic condi¬ 
tions, and McDonald and Jefferson (2008) suggested 
temperature limitations similar to those they found for 
Nothrotheriops shastensis. 


170 


Clyde Jones Memorial Volume 


Eptesicus fuscus (Big Brown Bat).—At least 
one specimen, not attributable to a zone, is of the large 
size typical of late Wisconsin big brown bats in the 
Southwest. Zones A and D also produced this species. 

f Panthera atrox (American Lion).—Although 
Smilodon cannot be entirely ruled out, measurements 
seem to better fit P. atrox. The single specimen is a 
distal right humerus retrieved, presumably from the 
cave surface, in 1966. 

Large cats likely were responsible for presence of 
large mammals such as horses. This species must have 
been an awesome predator, and even Puma concolor 
(Mountain Lion) can handle fairly large artiodactyls. 

Equus sp. (Horse).—Horse remains unidentifi¬ 
able to species are common from the surface to the 
lowest fossiliferous layer. Most are fragments of teeth 
or post-cranial elements. 

f Equus conversidens (Mexican Horse).—Sev¬ 
eral specimens are recognizable as this small Mexican 
Horse. Recognition primarily is by size, but a well 
preserved first phalanx has the typical proportions of 
this species. All but one specimen identified to this 
species were from disturbed sediments; the exception 
was from D/F (recorded as E in Table 1). 

f Hemiauchenia macrocephala (Big-headed 
Llama).—Four specimens are recognized as belonging 
to this camel; all are from disturbed sediments. The 
species is roughly the size of a Dromedary Camel, but 
with the proportions of a llama. It is difficult to envi¬ 
sion it reaching and entering the cave other than as parts 
carried in by predators or scavengers. As a cursorial 
animal, it seems best fit for relatively open country. It is 
widespread as a fossil in the southwestern Pleistocene. 

Odocoileus sp. (deer).—With the exception of 
remains from level A, specimens are labeled as from 
disturbed sites. A fresh apparent Mountain Lion kill 
was on the surface at the beginning of a field session. 

Antilocapra americana (Pronghorn).—Prong¬ 
horn remains were relatively rare (8 confident iden¬ 
tifications, 10 at the cf. level of certainty). Zones A, 
B, and F produced remains, but most specimens came 


from disturbed sediments. Pronghorn currently inhabit 
scrub-grassland on the flats above the cave. 

t Capromeryxfurcifer (Matthew’s Pronghorn).— 
Following White and Morgan (2011), Rancholabrean 
Capromeryx are considered to belong to C. furcifer. 
Some 25 elements of this small pronghorn are scattered 
through the site, zones A, B, D, and F. Remains are 
common in mid and late Wisconsin cave sites across 
southern New Mexico and into Chihuahua. A mixed 
diet and a habitat with clumps of shrubs and trees is 
suggested by several researchers (Bravo-Cuevas et ah 
2013). 

f Stockoceros conklingi (Conkling’s 
Pronghorn).—I follow Furlong (1943) in considering 
S. onusrosagris as a synonym of S. conklingi. This is a 
pronghorn intermediate in size between the diminutive 
C. furcifer and the large A. americana. Stockoceros 
conklingi remains are the most numerous of the larger 
mammals, with approximately 80 elements recovered 
and with all levels except E represented. 

From the common occurrence of numerous 
remains in caves, it is generally assumed that these 
antilocaprids used caves and shelters for protection 
against the elements. 

f Meleagris cf. M. crassipes (Big Foot Turkey).— 
A fragment of a tarsometatarsus bearing a spur too 
large for North American galliform birds other than 
Meleagris was recovered from zone “B (or possibly 
zone C)”. It is assumed from the labeling that the 
division between D and C was not clear. 

This turkey is somewhat smaller than M. gallopa- 
vo (Wild Turkey), thus likely represents M. crassipes. 
The specimen shows strong digestive corrosion. This 
species is known from several Guadalupe Mountains 
region faunas, including the mid Wisconsin fauna from 
Dry Cave (Rea 1980). 

f Coragyps occidentalis (Western Black Vul¬ 
ture).—Quite possibly ancestral to the Black Vulture 
(<Coragyps atratus ), this large form was the dominant 
vulture regionally, with the Turkey Vulture ( Cathartes 
aura) apparently absent from the Guadalupe Mountains 
region during the full glacial. Remains were found 


Harris—Pleistocene/Holocene Faunas from the Trans-Pecos 


171 


in zones A, B, C, and D, with a tentatively identified 
specimen from E. 

f Gymnogyps amplus (Pleistocene Condor).—The 
Pleistocene form of Gymnogyps was long considered 
as a separate species from the California Condor 
(G. californianus ), but later considered to be a large 
chronological subspecies of the modern California 
Condor. However, Syverson and Prothero (2010) pro¬ 
duced evidence that the Pleistocene form was indeed a 
separate species. From a practical viewpoint, much of 
the material from inland sites cannot as yet be parceled 
between the two species. New Mexico, Chihuahua, 
and Trans-Pecos Texas count 11 other Pleistocene sites 
containing Gymnogyps. It is identified from zone C 
and tentatively from zones B and F. 

f Geococcyx californianus conklingi (Conkling’s 
Roadrunner).—The distal one-fourth of a tarsometa- 
tarsus from Bl, level 4, is assignable to this large 
chronological subspecies of the Greater Roadrunner. 
It is considered a creature of the cooler summers of the 
late Wisconsin and may have survived into the early 
Holocene (Harris and Crews 1983). 


Corvus corax (Common Raven).—This is the 
common raven in the regional late Pleistocene record 
(Magish and Harris 1976) and at Sierra Diablo Cave. 
Zones B and C produced remains. Corvus corax oc¬ 
casionally occurs in the regional Chihuahuan Desert 
now, but is not common. 

Corax cryptoleucus (Chihuahuan Raven).—This 
is the current dominant species of Corvus in the vicin¬ 
ity of Sierra Diablo Cave. The Chihuahuan Raven is 
absent from the Guadalupe Mountains regional fossil 
record, though present in small numbers farther west 
in the Pendejo Cave mid and late Wisconsin (Harris 
2003). A single element is recognized from zone D. 

Pica hudsonicus (Black-billed Magpie).—A 
partial carpometacarpus from disturbed sediments 
was recovered. Magpies are common in northern New 
Mexico and are known to occasionally roam to the 
south. A single late Wisconsin element is recorded for 
the Guadalupe Mountains region. 


Discussion 


The question has been raised as to whether 
Fowlkes Cave adequately reflects late Pleistocene 
conditions south of the Guadalupe Mountains region 
(Harris 2016). The answer is that Fowlkes Cave does 
not—but also that data from Sierra Diablo Cave only 
partially clarifies the late Pleistocene faunal situation. 

Fowlkes Cave .—Dalquest and Stangl (1984) 
reported Holocene and Pleistocene mammalian faunas 
from Fowlkes Cave. Eater, anurans (Parmley 1988), 
snakes (Parmley 1990), a cotton rat (Stangl and Dal¬ 
quest 1991), and lizards (Parmley and Bahn 2012) were 
added to the Pleistocene fauna (Table 1). Bird remains 
have not been published. 

As interpreted by Dalquest and Stangl (1984), 
taxa unknown to coexist in the late Wisconsin faunas 
of the Guadalupe Mountains or elsewhere likely oc¬ 
curred within about 10 mi of Fowlkes Cave during 
the late Wisconsin. An example is the coexistence of 


desert taxa such as Dipodomys merriami, Onychomys 
arenarius, and Peromyscus eremicus together with 
non-desert species such as Sorex palustris, Sorex neo- 
mexicanus, and Marmola flaviventris. 

The cave is a sinkhole with the entrance well up 
a steep slope (Dalquest and Stangl 1984). The exact 
locality is unreported, but is about 10 km north of Kent, 
near the southern terminus of the relatively low (less 
than 1700 m) Apache Mountains. The elevation at the 
mouth of the cave was not given. As described by Dal¬ 
quest and Stangl (1984), the uppermost layer excavated 
consisted of about 30 cm of black silt. Below this layer 
was a sterile layer consisting of about 35 cm of rock 
fragments between 15 and 25 mm diameter (layer 1), 
then 20 cm of fragments between about 5 and 15 mm 
in diameter interspersed with fossils (layer 2), below 
which was a sterile layer of finer material about 30 cm 
thick (layer 3). 


172 


Clyde Jones Memorial Volume 


A partial lower jaw of an extinct miniature prong¬ 
horn ( Capromeryx ) was recovered in place in about the 
middle of layer 2, indicating a Pleistocene age. Mate¬ 
rial from that layer produced what Dal quest and Stangl 
(1984) thought to be a contemporaneous late Wisconsin 
fauna. Based largely on the size distribution of the fos¬ 
sil taxa, Dalquest and Stangl (1984) hypothesized that 
most of the fauna was introduced by Barn Owls (Tyto 
alba ) from within about 10 miles of the cave. 

The fauna (Table 1) includes both desert taxa 
and cool-adapted mesic forms. Dalquest and Stangl 
(1984:454) interpreted the taxa as contemporary and 
reconstructed the ecological conditions as follows: 

Cool streams with fringing borders of willows 
and meadowlands existed where there are now 
only dry washes, 10 km or more from the cave. 

The hillsides and aluvial [sic] fans supported 
somewhat more vegetation, including more 
junipers and perhaps some other trees. The 
creosote bush flats existed as today, but soils 
were sandy on terraces closer to the streams. 
Some of the typical desert mammals, such as 
Merriam kangaroo rat and long-tailed grass¬ 
hopper mouse, were just entering the area.... 

The scarcity of the Merriam kangaroo rat and 
long-tailed grasshopper mouse suggest that 
cooler climate of the late Pleistocene was just 
giving way to the hot, desert climate of modern 
conditions. 

The rational, aside from the occurrence of eco¬ 
logically mixed taxa in the same stratigraphic level, was 
that glaciation in the Guadalupe Mountains supplied 
cold, meltwater streams producing habitat for shrews 
and a vole, while Chihuahuan Desert habitat ruled the 
interfluvial areas. 

There are several problems with this scenario. 
For one, the Guadalupe Mountains were not glaciated, 
the nearest (and small) glacier occurring about 185 km 
to the NNW of the high peaks of the Guadalupe Moun¬ 
tains. Furthermore, drainage from north of the site 
originates from the southern end of the low Delaware 
Mountains, some 60 km south of the Guadalupes. The 
nearest rivers that would have carried glacial meltwater 


are the Pecos River some 80 km to the northeast and 
the Rio Grande about 90 km to the southwest. Also, 
regional reconstructions of vegetation based on packrat 
midden data indicates creosotebush rare or absent in 
the area until well into the Holocene, with woodland 
in the lowlands until at least 8,000 radiocarbon years 
before the present (Van Devender and Spaulding 1979). 

The dilemma facing Dalquest and Stangl (1984) 
is appreciated: the choice between a mixed late 
Wisconsin-Holocene fauna or, as suggested by their 
stratigraphic interpretation, a contemporaneous late 
Wisconsin fauna. They chose the latter. For reasons 
noted above and below, I take the opposite approach: 
that deposition started in the Pleistocene and continued 
well into the Holocene. 

The critical point revolves around the strati¬ 
graphic evidence. Dalquest and Stangl assumed the fos- 
siliferous layer was laid down within a relatively short 
period of time in the final stages of the late Wisconsin. 
However, layer 1 and the fossiliferous layer 2 consist 
of rock fragments and thus are necessarily lag deposits. 
Such deposits occur when fine particles are winnowed 
from them over time, concentrating relatively thick 
sedimentary layers into thinner layers of particles too 
large to be carried away by the available water flow. 
The alternative is a soilless, gravel-like surface layer 
surrounding the cave that nevertheless supported an 
abundant fauna. If, then, the fossiliferous layer repre¬ 
sents an appreciable length of time (evidenced in part by 
the relative fossil richness due to reduction of a thicker 
deposit), occurrence of late Wisconsin and Holocene 
taxa is explained. Absence of fossils in the overlying 
stratum may indicate relatively rapid deposition under 
severe Holocene climatic conditions. The black silt top 
deposits, containing a Holocene fauna (Dalquest and 
Stangl 1986) may well be of historic age, marking soil 
erosion due to the destruction of vegetation with the 
over-stocking of livestock. 

The stratigraphic evidence is upheld by the nature 
of the fauna. The mixture of taxa is not known from 
the extensive Wisconsin deposits of the Guadalupe 
Mountains region nor is it considered to be likely due 
to the more southern position of Fowlkes Cave. 


Harris—Pleistocene/Holocene Faunas from the Trans-Pecos 


173 


In summary: 

• Modern data indicate that such desert mammals 
as Merriam’s Kangaroo Rat occur under hot, dry 
climatic conditions; even where mesic conditions 
are nearby, such as along the Rio Grande, taxa 
typical of the Wisconsin Pleistocene are not 
found. Appeal to glacial meltwater doesn’t work. 

• Woodrat midden data indicate that hot, dry con¬ 
ditions did not arrive in the area until after the 
early Holocene. 

• Stratigraphic data strongly indicate that the 
lower fossiliferous stratum is a lag deposit mak¬ 
ing it feasible that deposition occurred over an 
extended span of time (late Pleistocene into mid 
or late Holocene), thus explaining the incompat¬ 
ible faunas. 

Sierra Diablo Cave. —The extensive disturbance 
of the upper portion of the cave sediments basically ren¬ 
ders much of its stratigraphic data useless. Especially in 
zone A, faunal evidence indicates extensive disturbance 
even where excavation data do not indicate such. This 
is true to a slightly lesser degree for the upper portion 
of zone B. A major drawback to interpretation is the 
rather obvious bias against recovery of the smaller 
faunal elements during the archaeological excavations. 

Aztlanolagus agilis was present about half way 
through stratum B and Desmodus stocki occurred near 
the base of that zone. These are thought to be markers 
of the mid Wisconsin and, as best as can be told, were 
in place stratigraphically. Taxa found in and below 
those levels are consistent with a mid Wisconsin age, 
though some taxa also would be expected to survive 
into the late Wisconsin and some into the Holocene. 

Potentially complicating matters, taxa west of 
the Guadalupe Mountains, such as at U-Bar Cave in 
the southwestern corner of New Mexico, show little 
faunal change between mid and late Wisconsin times 
as compared to those from Dry Cave (Harris 1989). 
It is possible that the late Wisconsin fauna at Sierra 
Diablo Cave, west of the Guadalupe-Delaware axis that 
presumably helps protect areas to the west from the full 
impact of Great Plains climate, differed from the mid 


Wisconsin fauna primarily by the loss of Aztlanolagus 
and Desmodus stocki. If so, with many of the same taxa 
present in both mid and late Wisconsin, there would be 
little in the Sierra Diablo Cave fauna that is unique to 
the late Wisconsin as opposed to the mid Wisconsin. 

Taxa that occur in the Guadalupe Mountains late 
Wisconsin that might indicate late (as opposed to mid) 
Wisconsin times if found at Sierra Diablo Cave include 
Pseudacris triseriata (Western Chorus Frog), Microtus 
longicaudus, Lepus townsendii (White-tailed Jackrab- 
bit), Sorex cinereus (Masked Shrew), Sorex palustris, 
and Mustela erminea (Ermine). The absence of these 
taxa from Sierra Diablo Cave, however, means little. 
They may be absent either because of the small sample 
size or because their geographic ranges did not reach 
south to Sierra Diablo Cave. Of these, only Sorex 
palustris is known to reach this far south (at Fowlkes 
Cave). 

In summary: 

• Strong disturbance of upper sediments prevents 
clear separation between deposits of different 
ages. 

• Deposition commenced in the mid Wisconsin 
and probably continued without interruption 
until the present. 

• There are no Pleistocene taxa present that would 
be expected to occur only in the late Wisconsin, 
as opposed to the mid Wisconsin. Absence of 
taxa typical of the Guadalupe Mountains late 
Wisconsin taxa, such are shrews, may be a result 
of sample size collecting bias, or may indicate 
real differences between the faunas. 

• Although the Sierra Diablo Cave fauna provides 
some information regarding the mid Wisconsin 
fauna in Trans-Pecos Texas south of the Guada¬ 
lupe Mountains region, characterization of the 
Pleistocene fauna must await discovery, excava¬ 
tion, and interpretation of a large fossil sample 
with firm stratigraphic evidence from the region. 


174 


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Acknowledgments 


I wish to thank Circle Ranch and Christopher 
Gill for allowing and encouraging excavations at Sierra 
Diablo Cave, as well as providing facilities for the field 
crews. David Carmichael and Javier Vasquez oversaw 
most of the excavations, and Angela Chavez was a driv¬ 
ing force in the early days. Financial support for the 
Vasquez excavations was from grants from the National 


Geographic Society and, through Vance T. Holiday, the 
Argonaut Archaeological Research Fund. Volunteer 
field crew members are too numerous to mention, but 
little could have been done without them. I also wish 
to thank Fred Stangl and Jim Goetze for reviewing the 
manuscript and making many useful comments. 


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Bravo-Cuevas, V. M., E. Jimenez-Hidalgo, M. A. Cabral-Perdo- 
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Furlong, E. L. 1943. The Pleistocene antelope, Stockoceros 
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Harris, A. H. 1970. Past climate of the Navajo Reservoir Dis¬ 
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Harris, A. H. 1987. Reconstruction of Mid-Wisconsin environ¬ 
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Harris, A. H. 1989. The New Mexican Late Wisconsin—East 
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Harris, A. H. 2003. The Pleistocene vertebrate fauna from 
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Harris, A. H. 2016. Pleistocene vertebrates of southwestern 
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Harris, A. H., and J. Hears! 2012. Late Wisconsin mammalian 
fauna from Dust Cave, Guadalupe Mountains National 
Park, Culberson County, Texas. Southwestern Naturalist 
57:202-206. 

Harris,A. H, and C. R. Crews. 1983. Conkling’s roadrunner—a 
subspecies of the California roadrunner? Southwestern 
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Kennedy, S., and C. Jones. 2006. Two new records of mammals 
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252:1-3. 

Logan, L. E. 1983. Paleoecological implications of the mam¬ 
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Logan, L. E., and C. C. Black. 1979. The Quaternary vertebrate 
fauna of Upper Sloth Cave, Guadalupe Mountains Na¬ 
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(H. H. Genoways and R. J. Baker, eds.). National Park 
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Lundelius, E. L., Jr. 1979. Post-Pleistocene mammals from 
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Magish, D. P, and A. H. Harris. 1976. Fossil ravens from the 
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Van Devender, T. R, K. B. Moodie, and A. H. Harris. 1976. 
The desert tortoise ( Gopherus agassizi ) in the Pleisto¬ 
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32:298-304. 

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Address of author: 

Arthur H. Harris 

University of Texas at El Paso Biodiversity Collections 
Departmen t of Biological Sciences 
The University of Texas at El Paso 
500 W University Ave. 

El Paso, TX 79968 
aharris @ utep. edu 


First Documented Record of Nutting’s Flycatcher (Myiarchus nuttingi ) for 

Texas 

Mark W. Lockwood 


Abstract 

A Nutting’s Flycatcher ( Myiarchus nuttingi ) was discovered near the mouth of 
Santa Elena Canyon, Big Bend National Park, Brewster County, Texas from 31 December 
2011 to 11 January 2012, providing a first record for the state. This species previously 
has been documented in the United States in Arizona and California. This paper discusses 
and provides context for this first documented and accepted state record for Texas. 

Key words: Brewster County, Myiarchus nuttingi , Nutting’s Flycatcher, state 
record, Texas 


Circumstances of the Occurrence 


On 31 December 2011, Brandon Percival and 
David Bradford discovered a Nutting’s Flycatcher 
(.Myiarchus nuttingi ) at the parking area near the 
mouth of Santa Elena Canyon in Big Bend National 
Park, Brewster County. Texas. The bird initially was 
heard calling from an area of dense mixed desert scrub 
bordering the parking area. This habitat is dominated 
by Honey Mesquite ( Prosopis glandulosa ), Salt Cedar 
(Tamarix sp.), Screwbean Mesquite ( Prosopis pu- 
bescens ), Roosevelt Weed (Baccharis neglecta ), and 
Gregg’s Acacia ( Acacia greggii ). Photographs were 
obtained and diagnostic vocalizations were heard. 
Recordings of the vocalizations were later obtained 


to fully document the occurrence. During the 12 days 
the bird was known to be present it appeared to remain 
in the vicinity of the initial discovery and was seen by 
many observers. It would remain in the dense vegeta¬ 
tion for long periods of time and then forage through 
the trees bordering the parking area. The bird was not 
particularly vocal, calling very irregularly and often for 
very short periods. It was not particularly wary and 
allowed close inspection when it was in view. Texas 
Bird Records Committee accepted the documentation 
(TBRC 2012-01; TPRF 2971), thus providing the first 
record for Texas (Carpenter 2013). 


Discussion 


Identification. —Virtually all of the species in 
the genus Myiarchus share very similar plumage char¬ 
acteristics that make identification of individual taxa 
more difficult. Nutting’s Flycatcher is very similar 
to the Ash-throated Flycatcher (M. cinerascens ) and 
these species were once considered to be conspecific 
(Lanyon 1961). The individual observed at Big Bend 
National Park exhibited plumage typical of Myiarchus 
flycatchers with a brownish-olive head contrasting with 
the gray-brown of the nape and back (Fig. 1). The au- 


riculars were lighter brown that the crown. The chin 
and breast were pale gray contrasting with a moderately 
bright yellow belly. The wings were darker brown with 
the coverts and tertials widely edged with tan. The 
primaries were widely edged with rufous. The tail was 
also dark brown with the inner webbing of all but the 
central rectricies mostly cinnamon with an obviously 
darker coloration on the outer web widening across 
the feather shaft towards the tip (Fig. 2). The bill was 
blackish with a flesh colored base to the lower man- 


177 


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Clyde Jones Memorial Volume 


Figure 1. This Nutting’s Flycatcher exhibited plumage characteristics consistent with populations in 
northwestern Mexico. This is the first accepted record for Texas. Photo by Mark W. Lockwood. 


Figure 2. The pattern of the underside of the retricies is an important plumage characteristic for separating 
Nutting’s Flycatcher (shown) from the similar Ash-throated Flycatcher. Photo by Mark W. Lockwood. 


Lockwood—First Record of Nutting’s Flycatcher for Texas 


179 


dible. The overall size was similar to an Ash-throated 
Flycatcher, but direct comparison was not made. The 
bird called very infrequently, but did give the distinc¬ 
tive, sharp wheeek! 

This plumage pattern of Nutting’s Flycatcher 
is shared with most of the other species in the genus 
Myiarchus. Of particular interest in the identification 
of this individual is Ash-throated Flycatcher, which is 
a common summer resident and uncommon and lo¬ 
cal winter resident along the Rio Grande in Brewster 
County. The Ash-throated Flycatcher differs very sub¬ 
tly from the Nutting’s and is best separated by voice. 
Ash-throated Flycatchers are very slightly larger and 
thinner bodied with a generally longer crest and larger 


bill. The undertail pattern of an adult Ash-throated 
Flycatcher differs in that the dark coloration of the 
outer webbing extends across the tip of the rectrices. 
However, first-winter Ash-throated have a tail pattern 
that is very similar to that of a Nutting’s. In general, 
Nutting’s Flycatchers are browner above with brighter 
yellow underparts. 

Distribution .—The Nutting’s Flycatcher ranges 
from northwestern Mexico southward to northwestern 
Costa Rica (Howell and Webb 1995, AOU 1998). 
The northernmost population of this species is found 
in northeastern Sonora within 75 km of the Arizona 
border (Howell et al. 2014) (Fig. 3). There has been 
a northward expansion of this species range during 


Figure 3. Range of Nutting’s Flycatcher and locations (indicated by black triangles) where the species has been 
documented in the United States. Range map based on Ridgely et al. (2003). 


180 


Clyde Jones Memorial Volume 


the past 50 years with the northernmost breeding area 
near Hermosillo in 1960 (Lanyon 1961) to an area in 
northeastern Sonora that is approximately 160 km far¬ 
ther north (Howell et al. 2014). In Sonora, Nutting’s 
Flycatchers are present year-round and are found in 
thorn forest habitats between sea level and 1800 m 
(Howell and Webb 1995). The first record for the 
United States was a bird collected near Roosevelt, 
Gila County, Arizona, on 8 January 1952 (Dickerman 
and Phillips 1953). Subsequent records for Arizona 
included one at Patagonia Lake State Park, Santa Cruz 
County, 14 December 1997 (Rosenberg 2001); one at 
Bill Williams Delta National Wildlife Refuge, Mohave 
County, 24 September 2008 (Stevenson and Rosenberg 
2009); and another was at the latter location in Decem¬ 
ber 201 l(Stevenson and Rosenberg 2012). Presumably 
this same individual was found on 30 November 2012 
at the Bill Williams Delta National Wildlife Refuge 
(Stevenson and Rosenberg 2013), which was followed 
by the first documented nesting record at the same lo¬ 
cation in April 2013 (Stevenson and Rosenberg 2014). 
This species has been found annually since, with six 


individuals present forming at least two pairs in the 
summer of 2014 (Stevenson and Rosenberg 2015). 
Interestingly, this location is approximately 580 km 
northwest of the northernmost population in Sonora, 
Mexico. There is also a single record for California, a 
single bird at Irvine, Orange County, from 11 November 
2000 through 26 March 2001 (California Bird Records 
Committee 2007). 

The Texas record may have been expected consid¬ 
ering the increase in numbers of Nutting’s Flycatchers 
in Arizona tied with the occurrence of other species 
of birds found primarily in western Mexico that have 
been documented in Texas. Species with similar ranges 
include Ruddy Ground-Dove ( Columbina talpacoti 
eluta). Rufous-backed Robin ( Turdus rufopalliatus ), 
Aztec Thrush ( Ridgwayia pinicola ), and Streak-backed 
Oriole ( Icterus pustulatus\ Lockwood and Freeman 
2014). The distance from known breeding areas to the 
location of the Texas record is approximately 480 km, 
which is closer than several of the Arizona records as 
well as the one from California. 


Acknowledgments 

Cliff Shackelford and Terry Maxwell reviewed helpful suggestions. Mark Stevenson provided infor- 

previous drafts of the manuscript and provided many mation about records from Arizona. 


Literature Cited 


California Bird Records Committee. 2007. Rare Birds of Cali¬ 
fornia (R. A. Hamilton, M. A. Patten, and R. A. Erickson, 
eds.). Western Field Ornithologists, Camarillo, CA. 

Carpenter, E. 2013. Texas Bird Records Committee Report 
for 2013. Bulletin of the Texas Ornithological Society 
46:52-56. 

Howell, S. N. G., I. Lewington, and W. Russell. 2014. Rare Birds 
of North America. Princeton University Press, Princeton. 

Howell, S. N. G., and S. Webb. 1995. A guide to the birds of 
Mexico and northern Central America. Oxford Univer¬ 
sity Press. 

Lanyon, W. E. 1961. Specific limits and distribution of the Ash- 
throated andNutting’sFlycatchers. Condor63:421-449. 

Lockwood, M. W., and B. Freeman. 2014. The TOS Handbook 
of Texas Birds. Texas A&M University Press, College 
Station. 


Ridgely, R. S., T. F. Allnutt, T. Brooks, D. K. McNicol, D. W. 
Mehlman, B. E. Young, and J. R. Zook. 2003. Digital 
distribution maps of the birds of the western hemisphere, 
version 1.0 NatureServe, Arlington, Virginia. 

Stevenson, M. M., and G. H. Rosenberg. 2009. Arizona: Fall 
migration. North American Birds 63:131-135. 

Stevenson, M. M., and G. H. Rosenberg. 2012. Arizona: Winter 
season. North American Birds 66:324-327. 

Stevenson, M. M., and G. H. Rosenberg. 2013. Arizona: Fall 
migration. North American Birds 67:131-135. 

Stevenson, M. M., and G. H. Rosenberg. 2014. Arizona: Spring 
migration. North American Birds 67:496M99. 


Lockwood—First Record of Nutting’s Flycatcher for Texas 


181 


Address of author: 

Mark W. Lockwood 

402 E. Harriet Ave. 

Alpine, Texas 79830 

mark, lockwood @ tpwd. texas. gov 


Bats of Kimball and Cheyenne Counties in the Panhandle of Nebraska 


Kenneth N. Geluso and Keith Geluso 


Abstract 

Despite numerous publications on bats throughout Nebraska, only a single record 
of a bat has been reported from Kimball County and none from Cheyenne County in 
the southwestern corner of the Nebraska panhandle. With concern about impacts of 
future wind-energy development on bat populations in the region, we conducted a study 
to examine the occurrence and seasonal activity of bats in those two counties. In 2010 
and 2011, we documented six species of bats in Kimball County—three migratory spe¬ 
cies ( Lasiurus cinereus, Lasiurus borealis, and Lasionycteris noctivagans) and three 
nonmigratory species (Eptesicus fuscus, Myotis ciliolabrum, and Myotis thysanodes). 
Only three species were documented in Cheyenne County— L. noctivagans, E. fuscus, 
and M. ciliolabrum. All six species were captured in the Pine Bluffs area of Kimball 
County, where pines, junipers, and rock outcrops were present; M. ciliolabrum was the 
only species captured in rocky, treeless areas of Cheyenne County; and L. noctivagans, 
E. fuscus, and M. ciliolabrum were captured or observed in riparian habitats in both 
counties. In 2010, a migratory wave of L. cinereus and L. borealis was documented in 
early August, and migration for both species seemed to be completed by the beginning 
of September. In contrast, migration of L. noctivagans seemed to begin and end later 
in the season. Migratory stopover sites in the Nebraska panhandle likely include the 
Pine Bluffs area as well as the Pine Ridge, Wildcat Hills, and Southern Wildcat Hills. 
Each stopover site provides an island of coniferous trees that can be used for resting and 
refuge, while also allowing a place for bats to refuel. In addition, waterways and other 
places with deciduous trees in the panhandle can be used as stopovers. Wind speeds 
are favorable for wind-energy development across the panhandle, including grassland 
areas. Construction of wind turbines in grasslands rather than in wooded areas in western 
Nebraska likely would reduce negative impacts on bat populations based on our capture 
success in the various habitats in the region. 

Key words: bats, distribution, Eptesicus fuscus, Lasionycteris noctivagans, La¬ 
siurus borealis, Lasiurus cinereus, migration, Myotis ciliolabrum, Myotis thysanodes, 
Nebraska, reproduction, seasonal activity, wind energy 


Introduction 


Despite a comprehensive review of Nebraska’s 
bats in 1979 (Czaplewski et al. 1979) and recent 
surveys of bats in western Nebraska (Benedict et al. 
2000; Benedict 2004; Geluso et al. 2004; Geluso et al. 
2013), only a single record of a bat has been reported 
from Kimball County (Silver-haired Bat, Lasionycteris 
noctivagans ; Geluso et al. 2004), whereas not a single 
species is known from Cheyenne County in the south¬ 


western corner of the Nebraska panhandle. However, 
on the basis of bat distributions and habitat availability 
in western Nebraska and neighboring states, as many as 
nine other species might occur in the region. Such spe¬ 
cies include the Fringed Myotis (. Myotis thysanodes), 
Long-legged Myotis ( Myotis volans ), Western Small¬ 
footed Myotis {Myotis ciliolabrum ), Big Brown Bat 
(Eptesicus fuscus). Eastern Red Bat (Lasiurus borealis). 


183 


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Clyde Jones Memorial Volume 


Hoary Bat ( Lasiurus cinereus ), Pallid Bat ( Antrozous 
pallidus), Townsend’s Big-eared Bat ( Corynorhinus 
townsendii ), and American Perimyotis ( Perimyotis 
subflavus ; Bogan and Cryan 2000; Geluso et al. 2005; 
Armstrong et al. 2011; Geluso et al. 2013). 

Construction of wind turbines for production of 
electrical energy has known adverse effects on bats 
(e.g., Kunz et al. 2007; Arnett et al. 2008). In the United 
States, wind-energy facilities are most detrimental to 
migratory species, especially L. noctivagans, L. ci¬ 
nereus, and L. borealis (Kunz et al. 2007; Grodsky et 
al. 2012). All three species inhabit western Nebraska 


(Czaplewski et al. 1979; Benedict 2004) and occur 
in the region during migration (Geluso et al. 2013). 
Understanding habitat use, relative abundance, and 
seasonality of migratory and nonmigratory species 
of bats in Kimball and Cheyenne counties is needed 
because of the potential for further development of 
wind power in the region. Thus far, seven operational 
wind turbines occur in the region (Nebraska Govern¬ 
ment Website 2016). Here we report our findings of 
a bat survey in those two counties during summer and 
early autumn, the time of year that most bat fatalities 
by wind turbines have been reported in North America 
(Kunz et al. 2007; Arnett et al. 2008). 


Study Area 


We conducted our study in Kimball and Chey¬ 
enne counties in the southwestern corner of Nebraska’s 
panhandle (Fig. 1). Study sites included coniferous 
woodlands in western Kimball County, pine-studded 
and treeless areas in Cheyenne County, and riparian 
habitats in both counties. Common and scientific names 
of plants mentioned below follow Kaul et al. (2006). 

Pine Bluffs area. —The Pine Bluffs area in 
western Kimball County is characterized by rolling 
hills with shallow canyons. The area occurs near 
the Nebraska border with Wyoming, southeast of the 
town of Pine Bluffs in Laramie County, Wyoming 
(Fig. 1). Hilltops and canyonsides contain scattered 
coniferous trees and occasional rock outcroppings. 
Common trees consist of Rocky Mountain Ponderosa 
Pine ( Pinus ponderosa ), Limber Pine ( Pinus flexilis ), 
and Rocky Mountain Juniper ( Juniperus scopulo- 
rum). Conspicuous shrubs in the area include Yucca 
(Yucca glauca ) and Fragrant Sumac ( Rhus aromatica ), 
whereas common grasses include gramas ( Bouteloua 
curtipendula, B. gracilis, and B. hirsuta ), Little Blue- 
stem ( Schizachyrium scoparium ), and Red Three-awn 
(Aristida purpurea). We sampled six water sources 
in this area. Two sites were situated on canyon floors 
(a metal stock tank 8.2 m in diameter [Fig. 2] and an 
earthen stock tank 44 by 18 m in length [Fig. 3]), and 
four sites were located in open, flat areas at different 
distances from canyonsides (an earthen stock tank 6.1 
by 1.5 m in length and metal stock tanks 5.5, 6.1, and 
9.1 m in diameter). 


The Pine Bluffs area only occupies about 6 km 2 
in Kimball County, but it is part of a larger wooded 
area that mostly occurs in southeastern Wyoming. 
The relatively rugged terrain in the Pine Bluffs area 
represents the northernmost part of a single, north-south 
lying escarpment that extends southward along the 
eastern border of Wyoming and terminates in northern 
Colorado. The total area in the tri-state region is about 
20 km 2 . 

Pine-studded area in Cheyenne County. —A 
unique habitat for Cheyenne County exists 4.7 km east 
of the town of Potter near a place named Point of Rocks 
(Fig. 1, localities 5-7). This area consists primarily 
of ponderosa pines and some junipers scattered on 
ridgetops and slopes of canyons (Fig. 4). Canyonsides 
also contain horizontal strips of rocky outcrops, yield¬ 
ing a terrace-like appearance. The total area containing 
trees is about 6 km 2 . We sampled a metal stock tank 
(3.0 m in diameter) in a grassy flat near the head of a 
canyon and placed a mist net over a large pond (70 
by 40 m in length) near a ranch house, 0.25 km from 
a canyonside. We also erected a net over land at the 
head of a canyon. 

Treeless region of southwestern Cheyenne 
County. —Most of southwestern Cheyenne County 
consists of gently rolling hills that are primarily used 
for grazing cattle; little agricultural land exists in the 
region compared to the remainder of the county. The 
landscape is essentially treeless except for occasional 


Geluso and Geluso—Bats of Kimball and Cheyenne Counties, Nebraska 


185 


Figure 1. Study sites in Kimball and Cheyenne counties in the southwestern corner of the Nebraska 
panhandle, where we attempted to capture bats in 2010 and 2011. Also shown are localities of speci¬ 
mens collected prior to our study. Localities in close proximity to each other are represented by the 
same circled number; for example, locality one represents five sites netted during our study and one 
site sampled in 2004 (Appendix). Black squares indicate six towns located along Lodgepole Creek. 


Figure 2. Over this metal stock tank (8.2 m in diameter) in the Pine Bluffs area of Kimball 
County, Nebraska, we captured five species of bats, including the only record of Fringed 
Myotis {Myotis thysanodes) from this county. 


186 


Clyde Jones Memorial Volume 


Figure 3. Over this earthen stock tank (44 by 18 m in length) in the Pine Bluffs area of 
Kimball County, Nebraska, we captured 32 Hoary Bats (Lasiurus cinereus ) and 18 Eastern 
Red Bats {Lasiurus borealis ) in a single evening (2 August—from dusk to dawn the fol¬ 
lowing morning), in what seemed to be a migratory wave in 2010. 


Figure 4. Area of scattered pines with some junipers near a place named Point of Rocks in 
Cheyenne County, Nebraska. Small maneuverable bats observed in this habitat probably 
were Western Small-footed Myotis {Myotis ciliolabrum). 


Geluso and Geluso—Bats of Kimball and Cheyenne Counties, Nebraska 


187 


human-planted windbreaks and trees growing around 
scattered ranch houses. Common grasses cover¬ 
ing the hills include gramas (same three species as 
above), Red Three-awn, Slender Wheatgrass ( Elymus 
trachycaulus ), Buffalo Grass (Buchloe dactyloides ), 
Needle-and-thread (Stipa comata ), and Japanese Brome 
(Bromus japonicus ). In addition, rocky outcrops break 
the landscape in many places. We sampled six water 
sources at different distances from those outcroppings 
(five metal stock tanks 1.8, 2.4, 6.1, 7.6, and 7.6 m in 
diameter and an earthen stock tank 25 by 8 m in length, 
Fig. 5). On two occasions, we placed a net across an 
open window and door of an abandoned rock house. 

Lodgepole Creek. —Lodgepole Creek is an east- 
west flowing, intermittent stream located in central 
Kimball and Cheyenne counties (Fig. 1). Lodgepole 


Creek is a tributary of the South Platte River, and it 
flows into the South Platte River near the town of Ovid, 
Sedgwick County, Colorado. Parts of the creek con¬ 
tain deciduous trees, whereas other parts are adjacent 
to grasslands or agricultural fields that lack trees. We 
sampled four localities along Lodgepole Creek from 
central Kimball County near the town of Kimball to 
eastern Cheyenne County near the town of Lodgepole 
(Fig. 1). Each site contained Plains Cottonwoods 
(Populus deltoides ) and willows ( Salix , Fig. 6). Ameri¬ 
can Elms ( Ulmus americana ) also were noted along the 
creek near towns of Potter and Lodgepole, and a few 
Russian-olives (Elaeagnus angustifolia) were observed 
at the site near Lodgepole. The creek varied in width 
from 2.7 to 15.2 m at our study sites and was <1 m in 
depth along most reaches. 


Figure 5. One of many rock outcrops in the treeless region of Cheyenne County, Nebraska. We 
captured Western Small-footed Myotis ( Myotis ciliolabrum ) at this site, when the dry earthen 
stock tank in the photograph contained water. 


188 


Clyde Jones Memorial Volume 


Figure 6. Lodgepole Creek between towns of Potter and Sidney in Cheyenne County, Nebraska. 
We captured Silver-haired Bats (Lasionycteris noctivagans). Big Brown Bats ( Eptes ic us fuse us ), 
and Western Small-footed Myotis ( Myotis ciliolabrum ) at this site. 


Methods and Materials 


To capture bats, we used mist nets (Avinet Inc., 
Dryden, NY, USA) ranging in length from 2.6 to 18 m 
and set them at ground level over water sources such 
as metal stock tanks, earthen stock tanks, and streams. 
On three occasions, nets were placed over land. We 
attempted to capture bats during 23 nights—21 nights in 
2010 (28-30 June; 27 July; 1,2,5, and 31 August; 1-6, 
9, 29, and 30 September; and 1 and 3-5 October) and 
two nights in 2011 (29 and 31 July). During our study, 
eight of 20 sites (40%) were sampled more than once 
(two to seven times), and on eight of 23 nights (35%), 
more than one site was sampled at the same time. 

Nets were set before evening twilight, and on 
18 of 23 nights (78%), they were kept up until dawn 
the next morning. For each individual captured, we 
recorded time of capture, species, sex, forearm length, 
body weight, reproductive condition (i.e., for females— 
pregnant, lactating, or not noticeably reproductive), 
and age (young of the year or adult). Females were 
recorded as lactating if milk could be expressed from 


mammary glands. An individual was considered a 
young of the year only if cartilaginous epiphyseal 
plates in finger joints were visible to the unaided eye 
when the wing was transilluminated (Anthony 1988). 
Using that trait, all bats could be aged through August, 
but beginning in September, it became difficult to age 
some individuals because cartilage was no longer vis¬ 
ible. Thus, beginning in September, some individuals 
recorded as adults might have been young of the year 
with ossified epiphyseal joints. 

When sampling bats during our survey, the first 
bats to appear at dusk were always small and highly 
maneuverable. Upon capture, each of those small bats 
was a M. ciliolabrum. Therefore, in cases where small, 
maneuverable bats were observed at dusk but not cap¬ 
tured, we reported the observation as “M ciliolabrum 
(observation only).” 

Most bats were released after processing at cap¬ 
ture sites, but some were kept as voucher specimens 


Geluso and Geluso—Bats of Kimball and Cheyenne Counties, Nebraska 


189 


and deposited in the natural history collection at the 
University of Nebraska State Museum in Lincoln 
(UNSM). Coordinates of localities were determined 
with a handheld Global Positioning System (GPS 72, 
Garmin International, Olathe, Kansas), using North 
American Datum 1983 (NAD 83). We also checked 
museum databases (The Mammal Networked Informa¬ 


tion System, using the VertNet portal) and collections 
at UNSM and at the University of Nebraska at Kearney 
and Omaha for unpublished records of bats from Kim¬ 
ball and Cheyenne counties. Specific locality descrip¬ 
tions for voucher specimens and for sites where bats 
were not captured are given in the Appendix. 


Results 


During 23 nights of netting in the southwestern 
corner of the Nebraska panhandle, we captured 115 
bats representing six species—L. cinereus, L. borea¬ 
lis, L. noctivagans, E. fuscus, M. ciliolabrum, and M. 
thysanodes (Table 1, Appendix). Bats were captured 
at four of seven sites in Kimball County (57% success) 
and at five of 13 sites in Cheyenne County (38% suc¬ 
cess). In addition, M. ciliolabrum was observed but 
not captured at two additional sites in Kimball County 
and at one site in Cheyenne County (Table 1). Dates 
of capture for each species are summarized in Table 
2. Reproductively active individuals were observed 
for E. fuscus (pregnant and lactating individuals and 
volant young), M. ciliolabrum (pregnant and lactating 
individuals), and L. cinereus (volant young; Table 3, 
Appendix). 

We captured bats in three of the four habitats 
sampled—the Pine Bluffs area of Kimball County, the 
treeless region of Cheyenne County, and Lodgepole 
Creek in both counties. In the Pine Bluffs area where 
pines, junipers, and rock outcrops were present, we 
documented all six species of bats captured in our study 
(Table 1). In treeless areas of Cheyenne County that 
contained rock outcroppings, we only documented M. 
ciliolabrum (Table 1). Two individuals were captured 
over an earthen tank next to a rock outcropping (Fig. 
5), and one was captured over a metal stock tank near 
an abandoned rock house that contained bat feces. 
Along Lodgepole Creek, we captured L. noctivagans, 
E. fuscus, and M. ciliolabrum among deciduous trees 
in Cheyenne County, and along that creek in Kimball 
County, we captured L. noctivagans and E. fuscus and 
observed M. ciliolabrum (Table 1). In the pine-studded 


area of Cheyenne County that contained rocky out¬ 
crops, we were unable to capture any bats but suspected 
that small, maneuverable bats observed at dusk were 
M. ciliolabrum (Table 1). 

We captured bats from late June (28th) to early 
October (5th). Most individuals of L. cinereus (36 of 
37) and all those ofL. borealis {19 of 19) were captured 
in late July and early August 2010 (Table 2). Each was 
captured in the Pine Bluffs area either over a large, 
unobstructed earthen tank (two Hoary Bats and one 
Eastern Red Bat in July; 32 Hoary Bats and 18 Eastern 
Red Bats in August, Fig. 3) or over a nearby metal tank 
(two Hoary Bats in August, Fig. 2). Another L. cinereus 
was captured over the metal tank on 31 August. The 
earthen tank was dry on subsequent visits in 2010 and 
2011. Our first captures of L. noctivagans occurred 
in early September in both Kimball and Cheyenne 
counties, and later captures continued through late 
September and early October, respectively (Table 2). 
For nonmigratory species, E. fuscus was captured from 
late June to early September, M. ciliolabrum from late 
June to late September, and our only capture of M. 
thysanodes occurred on 29 September (Table 2). 

Besides our captures and observations of M. cili¬ 
olabrum, we discovered four unpublished records of 
that species from Kimball County—three from the Pine 
Bluffs area and one from near the town of Kimball. In 
the Pine Bluff area, one female each was captured on 
15 July, 17 July (lactating), and 4 August 2004; and 
near Kimball, a male was captured on 16 May 1964 
(Appendix). 


190 


Clyde Jones Memorial Volume 


Table 1. Number of bats captured at seven sites in Kimball County and 13 sites in Cheyenne County, Nebraska, in 
2010 and 2011. Numbers in parentheses correspond to those in Figure 1. Detailed locality descriptions for each site 
are given in the Appendix. 


Site description 

Lasiurus 

cinereus 

Lasiurus 

borealis 

Lasionycteris 

noctivagans 

Eptesicus 

fuscus 

Myotis 

ciliolabrum 

Myotis 

thysanodes 

Pine Bluffs area, Kimball Co. 

(1) metal tank 

- 

- 

- 

- 

- 

- 

(1) small earthen tank 

- 

- 

- 

1 

1 

- 

(1) large earthen tank 

34 

19 

- 

20 

3 

- 

(1) metal tank 

- 

- 

- 

- 

__i 

- 

(1) metal tank 

3 

- 

1 

1 

5 

1 

(2) metal tank 

- 

- 

- 

- 

- 

- 

Pine-studded area, Cheyenne Co. 

(5) large pond 

- 

- 

- 

- 

- 

- 

(6) over land 

- 

- 

- 

- 

__i 

- 

(7) metal tank 

- 

- 

- 

- 

- 

- 

Treeless region, Cheyenne Co. 

(9) metal tank 

- 

- 

- 

- 

- 

- 

(9) metal tank 

- 

- 

- 

- 

1 

- 

(9) next to rock house 

- 

- 

- 

- 

- 

- 

(10) metal tank 

- 

- 

- 

- 

- 

- 

(11) metal tank 

- 

- 

- 

- 

- 

- 

(11) metal tank 

- 

- 

- 

- 

- 

- 

(11) earthen tank 

- 

- 

- 

- 

2 

- 

Lodgepole Creek, Kimball Co. 

(4) at Kimball 

- 

- 

1 

1 


- 

Lodgepole Creek, Cheyenne Co. 

(8) near Potter 

- 

- 

2 

- 

- 

- 

(12) between Potter & Sidney 

- 

- 

4 

11 

2 

- 

(13) near Lodgepole 

- 

- 

2 

- 

- 

- 

Total captured 

37 

19 

10 

34 

14 

1 


Observation only; see Methods and Materials 


Geluso and Geluso—Bats of Kimball and Cheyenne Counties, Nebraska 


191 


Table 2. Dates of capture for bats in Kimball and Cheyenne counties, Nebraska, in 2010 and 2011. Dates also include 
probable observations of Myotis ciliolabrum. For each date, p.m. refers to individuals captured from dusk to midnight, 
and a.m. refers to those captured after midnight to dawn. All dates refer to 2010, except 29 July, 31 July, and 1 August 
(a.m.), which refer to 2011. 


Date 

Lasiurus 

cinereus 

Lasiurus 

borealis 

Lasionycteris 

noctivagans 

Eptesicus 

fuscus 

Myotis 

ciliolabrum 

Myotis 

thysanodes 

28 June (p.m.) 

- 

- 

- 

1 

- 

- 

29 June (a.m.) 

- 

- 

- 

-- 

1 

- 

30 June (p.m.) 

- 

- 

- 

6 

1 

- 

1 July (a.m.) 

- 

- 

- 

3 

-- 

- 

27 July (p.m.) 

1 

1 

- 

8 

4 

- 

28 July (a.m.) 

1 

-- 

- 

3 

1 

- 

29 July (p.m.) 

-- 

-- 

- 

-- 

__i 

- 

31 July (p.m.) 

-- 

-- 

- 

-- 

__i 

- 

1 August (a.m.) 

-- 

-- 

- 

1 

-- 

- 

1 August (p.m.) 

-- 

-- 

- 

-- 

2 

- 

2 August (p.m.) 

6 

8 

- 

8 

2 

- 

3 August (a.m.) 

28 

10 

- 

1 

-- 

- 

31 August (p.m.) 

1 

-- 

- 

-- 

__i 

- 

3 September (p.m.) 

-- 

-- 

1 

2 

__i 

- 

4 September (p.m.) 

-- 

-- 

1 

1 

__i 

- 

5 September (p.m.) 

-- 

-- 

-- 

-- 

__i 

- 

6 September (p.m.) 

-- 

-- 

-- 

-- 

__i 

- 

7 September (a.m.) 

-- 

-- 

2 

-- 

-- 

- 

29 September (p.m.) 

-- 

-- 

-- 

-- 

1 

1 

30 September (a.m.) 

-- 

-- 

1 

-- 

-- 

-- 

30 September (p.m.) 

-- 

-- 

-- 

-- 

2 

-- 

1 October (a.m.) 

-- 

-- 

1 

-- 

-- 

-- 

1 October (p.m.) 

-- 

-- 

2 

-- 

-- 

-- 

4 October (p.m.) 

-- 

-- 

-- 

-- 

__i 

-- 

5 October (p.m.) 

-- 

-- 

2 

-- 

-- 

-- 

Total captured 

37 

19 

10 

34 

14 

1 


Observation only; see Methods and Materials 


192 


Clyde Jones Memorial Volume 


Table 3. Dates of reproductive activity for bats in Kimball and Cheyenne counties, Nebraska, in 
2010. None of the dates for lactation extend the period of lactation for a species in the state, and 
none of the dates for juveniles provide earlier dates when volant young first appear in Nebraska 
or extend dates in the state when young still can be recognized by cartilage in their finger joints. 


Species 

June 

July 

August 

September 

Eptesicus fuscus 

Pregnant 

30 1 

1 

- 

- 

Lactating 

30 

27, 28 

2 

- 

Volant young 

- 

- 

2,3 

3 

Myotis ciliolabrum 

Pregnant 

30 2 

- 

- 

- 

Lactating 

- 

27 

1,2 

- 

Lasiurus cinereus 

Volant young 

- 

27 

2,3 

- 


female contained a single fetus with a crown-to-rump length of 22 mm. 
2 Female contained a single fetus with a crown-to-rump length of 17 mm. 


Discussion 


At least six species of bats occur in the south¬ 
western corner of the Nebraska panhandle—three 
migratory species (L. cinereus, L. borealis, and L. noc- 
tivagans) and three nonmigratory species (. E.Juscus, M. 
ciliolabrum, and M. thysanodes ). The only previously 
published record from this region is an L. noctivagans 
that was discovered dead in Kimball County in 1988 
(Geluso et al. 2004; Appendix). Thus, our captures of 

L. cinereus, L. borealis, E.Juscus, M. ciliolabrum, and 

M. thysanodes represent new records of occurrence 
for Kimball County. We only captured three species 
in Cheyenne County (L. noctivagans, E. fuscus, and 
M. ciliolabrum ), and each represents a new record of 
occurrence for that county. For each newly reported 
species in Kimball County, the closest locality of oc¬ 
currence in Nebraska is from the Southern Wildcat 
Hills in Banner County, 41 km north for L. cinereus, 
38 km northwest for E. fuscus, and 39 km north for the 
other species (Geluso et al. 2013). For new records in 
Cheyenne County, the closest record in Nebraska for L. 
noctivagans is 45 km north-northeast in Morrill County 
and for E. fuscus and M. ciliolabrum, 46 km north in 
Morrill County (Geluso et al. 2013). Because of a lack 


of historical sampling of bats in the southwestern comer 
of the panhandle (Jones 1964; Czaplewski et al. 1979; 
Benedict et al. 2000; Benedict 2004), records reported 
herein from Kimball and Cheyenne counties are best 
referred to as range extensions and not expansions in 
distribution (sensu Frey 2009). 

The Pine Bluffs area had the greatest species 
richness of the four habitats sampled during our study 
(six species, Table 1), and each of those species also is 
known from the Southern Wildcat Hills, 37 km north of 
the Pine Bluffs area (Geluso et al. 2013). Similar to the 
habitat in the Pine Bluffs area, hilltops and canyonsides 
of the Southern Wildcat Hills contained ponderosa 
pines, junipers, and rock outcroppings. However, 
conifers were denser, canyons were deeper, and rocks 
were more abundant in the Southern Wildcat Hills. In 
that wooded and rugged terrain, Geluso et al. (2013) 
suspected that three additional species might reside 
in those hills, as we did for the Pine Bluffs area (i.e., 
M. volans, C. townsendii, and A. pallidus). In both 
areas, however, none of those species were captured in 
either survey. Geluso et al. (2013) discussed possible 


Geluso and Geluso—Bats of Kimball and Cheyenne Counties, Nebraska 


193 


reasons for their seeming absence (e.g., low population 
densities, lack of certain habitat requirements, or both), 
which also apply to the Pine Bluffs area. 

Our only capture of M. thysanodes occurred in 
the Pine Bluffs area of Kimball County. In Nebraska, 
M. thysanodes is a Tier 1 species of concern due to its 
limited distribution and rarity in the Wildcat Hills and 
Pine Ridge regions of the state (Schneider et al. 2011). 
Our capture in Kimball County, along with recent cap¬ 
tures in the Southern Wildcat Hills (Banner County, 
Geluso et al. 2013), demonstrates that the species has a 
greater distribution in the state than previously known. 
In Nebraska thus far, M. thysanodes only occurs in ar¬ 
eas associated with coniferous woodlands and forests 
(Czaplewski et al. 1979; Geluso et al. 2013; this study). 
Our capture on 29 September and another individual 
captured on that date in the Wildcat Hills (Geluso et 
al. 2013) represent the latest seasonal date for M. thy¬ 
sanodes in Nebraska (Table 2). Although no winter 
records of M. thysanodes are known from Nebraska, 
hibernating individuals have been reported near the 
Nebraska panhandle in southwestern South Dakota and 
southeastern Wyoming (Turner and Jones 1968; Martin 
and Hawks 1972; Bogan and Cryan 2000; Geluso et al. 
2005). Caves and mines in neighboring states might be 
important shelters for hibernating species of bats that 
spend summer months in the panhandle of Nebraska. 
This species and others likely also hibernate within 
Nebraska in rock fissures or other suitable hibernacula. 
Information on wintering habits for all species of bats 
that overwinter in the state is warranted because white- 
nose syndrome continues to spread westward (Frick et 
al. 2010), and recently, the fungus associated with the 
syndrome has been documented in eastern Nebraska 
(WNS 2015). 

Myotis ciliolabrum was the only species captured 
in the treeless region of southwestern Cheyenne Coun¬ 
ty, and although we did not capture a single bat in the 
pine-studded area of that county, small bats observed 
in the canyons likely were that species (Table 1). A 
common feature in both habitats was the presence of 
numerous rock outcroppings. Throughout most of its 
distribution, presence of rocks seems to be an impor¬ 
tant habitat requirement for M. ciliolabrum (Jones et 
al. 1983), and our captures and observations in south¬ 
western Cheyenne County support that conclusion. 


Besides M. ciliolabrum , we doubt that other species of 
bats commonly occur in those habitats of the county. 

Although we only captured three species in ripar¬ 
ian habitats along Lodgepole Creek (L. noctivagans, 
E. fuscus, and M. ciliolabrum , Table 1), we suspect 
that L. cinereus, L. borealis , and possibly other species 
also periodically inhabit that wooded waterway. In the 
panhandle, for example, L. cinereus has been captured 
in cottonwood communities along Pumpkin Creek and 
the North Platte River (Geluso et al. 2013), and L. bo¬ 
realis is known from the town of Bridgeport near the 
North Platte River (Benedict et al. 2000). P. subflavus 
is another species that likely occurs in riparian habitats 
in Kimball and Cheyenne counties. During summer in 
the western part of its distribution, P. subflavus roosts 
in areas with isolated deciduous trees surrounded by 
grasslands or deserts (Geluso et al. 2005; White et al. 
2006; Valdez et al. 2009). This species has expanded 
its distribution across the Great Plains in recent decades 
(Geluso et al. 2005), and records are now available 
from northeastern Colorado and southeastern Wyoming 
(Bogan and Cryan 2000; Armstrong et al. 2006). Lack 
of captures ofP subflavus during our study might be re¬ 
lated to (1) minimal netting done by us along the creek, 
(2) a low population density in the area, (3) difficulty 
in capturing individuals in mist nets (e.g., White et al. 
2016), or (4) some combination of the above. 

Migration and migratory stopover sites. —During 
our study in the southwestern corner of the panhandle, 
autumn migration of L. cinereus and L. borealis seemed 
to have begun in late July/early August (Table 2), a 
pattern similar to that observed in more northern areas 
of the panhandle (Benedict 2004; Geluso et al. 2013). 
Our captures of 34 Hoary Bats and 18 Eastern Red Bats 
on a single August night likely represent a migratory 
wave through the Pine Bluffs area. On that night, bats 
were captured over two nearby stock tanks, but six days 
prior in late July, only two Hoary Bats and one Eastern 
Red Bat were captured from dusk to dawn at the same 
two sites (Table 2). Moreover, 73% of total captures 
in early August (28 of 34 Hoary Bats and 10 of 18 
Eastern Red Bats) occurred after midnight (0004-0448 
h), which further suggests the presence of migratory 
individuals because in temperate climates, relatively 
few insectivorous bats are captured over water after 
midnight; on typical summer nights, most individuals 


194 


Clyde Jones Memorial Volume 


are captured < 3 h after sunset (e.g., Jones 1965). In 
addition, not one female of either species was lactating 
that night (n = 26 adults), indicating a cohort of females 
free to travel without nursing nonvolant young. Sex 
ratios of adult migrants were nearly 1:1 for L. cinereus 
(14 females and 16 males) and 2:1 for L. borealis (12 fe¬ 
males and 6 males). The migratory wave also included 
four young Hoary Bats (three males and one female), 
three of which were captured after midnight (female 
at 0131, male at 0222, and male at 0416 h). Migra¬ 
tory young with and without visible cartilage in their 
finger joints also have been reported in early August 
in New Mexico (Findley and Jones 1964; Geluso and 
Geluso 2004). Findley and Jones (1964) stated that 
different age groups of L. cinereus might migrate at 
different times during southward migration; however, 
our captures in the Pine Bluffs area suggest that young 
and adults sometimes travel together. Although autumn 
migration of L. cinereus and L. borealis seemed to be 
completed by the beginning of September in the Pine 
Bluffs area (Table 2), migratory individuals have been 
captured as late as 15 September for Hoary Bats and 
as late as 29 September for Eastern Red Bats in other 
parts of the Nebraska panhandle (Geluso et al. 2013). 

In the study area, autumn migration of L. noctiva- 
gans began at least a month later than that of Hoary and 
Eastern Red bats (Table 2). For example, we captured 
our first Silver-haired Bats in early September but did 
not capture a single individual in early August, when 
many Hoary and Eastern Red bats passed through 
the area. Similar to those findings, most migratory 
Silver-haired Bats in the Wildcat Hills and surround¬ 
ing area also were captured in September, with only a 
single capture in late August (Geluso et al. 2013). In 
addition, migratory waves of L. noctivagans have been 
reported in late August and mid September in Ontario, 
Canada (McGuire et al. 2012). In western Nebraska, 
autumn migration also ends later for L. noctivagans. 
For example, Silver-haired Bats have been captured on 
the wing as late as 4 November, whereas late captures 
for L. cinereus and L. borealis do not extend beyond 
September in western Nebraska (Geluso etal. 2013). In 
Kimball and Cheyenne counties, half of our captures of 
L. noctivagans occurred in October (Table 2). Similar 
to the sex ratio of migrant Hoary Bats, the ratio for 
Silver-haired Bats was 1:1 (5 males and 5 females), 
with some individuals possibly being young of the 


year with recent epiphyseal ossification (see Methods). 
Geluso et al. (2013) reported that in the Wildcat Hills 
and surrounding area, Silver-haired Bats occurred in 
and around coniferous forests only during spring and 
autumn migration; however, we discovered that they 
also use waterways lined with deciduous trees, at least 
during autumn migration. For example, nine of 10 
captures of L. noctivagans occurred along Lodgepole 
Creek, and only one capture occurred in the Pine Bluffs 
area (Table 1). 

All three migratory species were captured during 
autumn migration in the Pine Bluffs area of western 
Kimball County (Tables 1 and 2). Similarly, those spe¬ 
cies have been documented during autumn migration 
in other coniferous habitats to the north, including the 
Pine Ridge (Sioux, Dawes, and Sheridan counties), 
Wildcat Hills (Scotts Bluff, Banner, and Morrill coun¬ 
ties), and Southern Wildcat Hills (Banner and Morrill 
counties, Geluso et al. 2004; Geluso et al. 2013). Those 
four areas in the panhandle of Nebraska likely provide 
important stopover sites for migratory bats travelling 
through a region that is dominated by grasslands. 
Each area provides an island of trees that can be used 
for resting and refuge, while also allowing a place for 
bats to refuel (see McGuire et al. 2012). In addition, 
waterways and other places with deciduous trees in the 
panhandle can be used as stopovers during migration 
(e.g., L. noctivagans. Table 1), because each migratory 
species ( L. cinereus, L. borealis, and L. noctivagans) is 
known to roost in both coniferous and deciduous trees 
(Ammerman et al. 2012). 

Bats and development of wind power in Ne¬ 
braska. —According to the American Wind Energy 
Association, Nebraska is the fourth windiest state in 
the United States and ranks sixth in the nation for the 
greatest potential to generate electricity from wind 
power (AWEA2007; Power Online 2010). As of2014, 
Nebraska has 473 operational wind turbines in the state, 
seven of which are located in a grassland area of Kim¬ 
ball County (4.6 km N, 3.1 km W Kimball; Nebraska 
Government Website 2016). Not surprisingly, efforts 
are slated to expand the development of wind energy 
in Nebraska (Nebraska Legislature 2009). 

Observations of bat and avian fatalities beneath 
wind turbines indicate that wind energy is not as en- 


Geluso and Geluso—Bats of Kimball and Cheyenne Counties, Nebraska 


195 


vironmentally friendly as once thought (Kunz et al. 
2007; Arnett et al. 2008). In the eastern United States, 
large numbers of bats, mainly L. borealis, L. cinereus, 
and L. noctivagans, have been killed at utility-scale, 
wind-energy facilities, especially by turbines situated 
along forested ridgetops (Kunz et al. 2007). Although 
wind turbines frequently kill species that migrate long 
distances, fatalities also have been reported for more 
sedentary species such as E. fuscus, P. subflavus, M. 
lucifugus, and M. septentrionalis (Kunz et al. 2007; 
Grodsky et al. 2012). Thus, development of wind 
power could impact populations of migratory (L. 
cinereus, L. borealis, and L. noctivagans) and nonmi- 
gratory (E. fuscus, M. ciliolabrum, andM thysanodes ) 
species in western Nebraska, if wind-energy facilities 
were constructed in areas frequented by those species. 

Unfortunately for the bats, probable stopover sites 
for many migratory individuals as well as areas used by 
some nonmigratory species in the panhandle of Nebras¬ 
ka (i.e., Pine Ridge, Wildcat Hills, Southern Wildcat 
Hills, and Pine Bluffs area) seem to include ideal loca¬ 
tions for generating electricity from wind power. Not 
only are ridgetops and escarpments available, but wind 
speeds also are ideal on the basis of average speeds 80 
m above the ground (NREL 2010; commercial wind 


turbines range in height from 45 to 100 m, Kunz et 
al. 2007). Areas with average wind speeds > 6.5 m/s 
at an 80-m height are generally considered to have a 
wind resource suitable for wind-energy development 
(Windexchange 2015), and almost the entire panhandle 
of Nebraska has wind speeds > 7.5 m/s 80 m above 
the ground (NREL 2010). Highest wind speeds in the 
panhandle occur in the Wildcat Hills of Banner County 
and Pine Bluffs area of Kimball County (> 10.0 m/s). 
Wind speeds also are favorable in the Pine Ridge and in 
grassland areas of the panhandle (NREL 2010). On the 
basis of captures of migratory and nonmigratory species 
of bats in our study (Table 1) and in other areas of the 
panhandle of Nebraska (Benedict et al. 2000; Benedict 
2004; Geluso et al. 2004; Geluso et al. 2013), it seems 
that construction of wind turbines in open grassland 
areas would reduce negative impacts on bat popula¬ 
tions. After acoustically monitoring bat activity across 
different landscapes, Baerwald and Barclay (2009) 
also concluded that bat fatalities probably would be 
minimized by building wind-energy facilities in prairies 
rather than in wooded areas that provide stopover sites 
for migrating tree bats. To assess potential impacts of 
utility-scale, wind-energy facilities on bat populations, 
it is necessary to know what species occur and when 
they occur in areas that might support such facilities. 


Acknowledgments 


We thank various landowners in the region for ac¬ 
cess to their lands. We also thank Claire Dodd, Kelley 
Penney, David Sempek, and Jeremy White (University 
of Nebraska at Omaha, UNO) for assistance in the 
field; Thomas Labedz (University of Nebraska State 
Museum) for assistance with museum matters; Angie 
Fox (Scientific Illustrator, University of Nebraska State 


Museum) for preparing Figure 1; and Jeffrey Hueb- 
schman (University of Wisconsin—Platteville) and 
Jeremy White (UNO) for constructive comments on 
an earlier version of this manuscript. This project was 
funded by the Nebraska Game and Parks Commission 
(Lincoln, Nebraska). 


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Addresses of authors: 

Kenneth N. Geluso 

Department of Biology 
University of Nebraska at Omaha 
Omaha, NE 68182 
kgeluso @ unomaha. edu 


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Keith Geluso 

Department of Biology 
University of Nebraska at Kearney 
Kearney, NE 68849 
gelusokl @ unk. edu 


198 


Clyde Jones Memorial Volume 


Appendix 

Localities where we attempted to capture bats in Kimball and Cheyenne counties, Nebraska, in 2010 and 2011. 
Numbers in parentheses before each locality correspond to circled numbers in Figure 1. Localities in close prox¬ 
imity to each other are represented by the same number. Listing of sites follows the order shown in Table 1. A 
description of the netting site, total number of bats captured, and dates of netting are given after each locality. 
Dates also include probable observations of Myotis ciliolabrum. For each date, p.m. refers to individuals captured 
from dusk to midnight, and a.m. refers to those captured after midnight to dawn. In addition, the number of males 
and females captured of each species is given with corresponding information on reproductive condition and age. 
Voucher specimens are shown in brackets with their museum number at the University of Nebraska State Museum 
in Lincoln (UNSM) and University of Wyoming, Museum of Vertebrates in Laramie (UWYMV). At the end of 
the appendix, we also provide information for specimens collected prior to our study. 


Pine Bluffs area in Kimball County 

(1) 2.79 km S, 0.51 km W Intersection of Stateline Road and Interstate 1-80, 41°09.441'N, 104°03.012'W, metal 
tank 6.1 m in diameter in open, flat area near canyonside, no bats captured on night of 31 July 2011, including 
after midnight. 

(1) 2.8 km S, 0.5 km W Intersection of Stateline Road and Interstate 1-80, 41°09.437'N, 104°03.002'W, small 
earthen tank 6.1 by 1.5 m in length in open, flat area near canyonside, two bats captured: 28 June 2010, p.m.— E. 
fuscus (one adult male [UNSM 30084]); 29 June 2010, a.m.— M. ciliolabrum (one adult male [UNSM 30091]); 
31 August 2010—none captured including after midnight. 

(1) 2.6 km S, 0.1 km W Intersection of Stateline Road and Interstate 1-80, 41°09.555'N, 104°02.759'W, large 
earthen tank 44 by 18 m in length on canyon floor, 76 bats captured: 27 July 2010, p.m.— E. fuscus (six lactat- 
ing females, two adult males), M. ciliolabrum (one lactating female), L. cinereus (one young-of-the-year female 
[UNSM 30066]), andL. borealis (one adult female [UNSM 30071]); 28 July 2010, a.m.— E. fuscus (one lactating 
female, one adult female, one adult male), M. ciliolabrum (one adult male), and L. cinereus (one adult female); 

2 August 2010, p.m.— E. fuscus (three lactating females, three adult females, one adult male, one young-of-the- 
year male), M. ciliolabrum (one lactating female), L. cinereus (two adult females, three adult males, one young- 
of-the-year male), and L. borealis (four adult females, four adult males); 3 August 2010, a.m.— E. fuscus (one 
young-of-the-year male), L. cinereus (10 adult females, 13 adult males [UNSM 30068], one young-of-the-year 
female, two young-of-the-year males), and L. borealis (eight adult females, two adult males). 

(1) 3.2 km S, 0.1 km W Intersection of Stateline Road and Interstate 1-80,41°09.235'N, 104°02.737'W, metal tank 
5.5 m in diameter in open, flat area near canyonside, no bats captured on night of 5 September 2010, including 
after midnight; Myotis ciliolabrum observed p.m. 

(1) 2.5 km S, 0.1 km E Intersection of Stateline Road and Interstate 1-80, 41°09.607'N, 104°02.622'W, metal 
tank 8.2 m in diameter on canyon floor, 11 bats captured: 27 July 2010, p.m.— M. ciliolabrum (one adult female 
[UNSM 30094], one lactating female, one adult male); 2 August 2010, p.m.— M. ciliolabrum (one adult female); 

3 August 2010, a.m.— L. cinereus (two adult females); 31 August 2010, p.m.— L. cinereus (one adult male) and 
Myotis ciliolabrum (observation only); 29 September 2010, p.m.— M. ciliolabrum (one adult male) and M. thy- 
sanodes (one adult male [UNSM 30090]); 30 September 2010, a.m.— L. noctivagans (one adult female [UNSM 
30079]); 4 October 2010, p.m.— Myotis ciliolabrum (observation only); 29 July 2011, p.m.— Myotis ciliolabrum 
(observation only); 31 July 2011, p.m.— Myotis ciliolabrum (observation only); 1 August 2011, a.m.— E. fuscus 
(one adult male [UNSM 30097]). 


Geluso and Geluso—Bats of Kimball and Cheyenne Counties, Nebraska 


199 


(2) 0.8 km S, 1.5 km E Intersection of Stateline Road and Interstate 1-80, 41°10.568'N, 104°01.593'W, metal 
tank 9.1 m in diameter in open, flat area near canyonside, no bats captured on night of 3 October 2010, including 
after midnight. 

Pine-studded area in Cheyenne County 

(5) 0.5 km S, 4.4 km E Potter, 41°12.773'N, 103°15.786'W, large pond 70 by 40 m in length, 0.25 km from a 
canyonside, no bats captured on night of 29 June 2010, including after midnight. 

(6) 0.1 kmN, 4.7 km E Potter, 41°13.142'N, 103°15.514'W, over land at canyon head, no bats captured on night 
of 1 August 2010 (net removed before midnight); Myotis ciliolabrum observed. 

(7) 5 km E Potter, 41°13.022'N, 103°15.342'W, metal tank 3.0 m in diameter in flat area at canyon head, no bats 
captured on night of 1 August 2010 (net removed before midnight); no bats captured on night of 5 August 2010, 
including after midnight. 

Treeless region in Cheyenne County 

(9) 16.3 km S, 0.5 kmE Potter, 41°04.239'N, 103°18.366'W, metal tank 7.6 m in diameter in rolling hills, no bats 
captured on night of 30 September 2010, including after midnight. 

(9) 16.2 km S, 1.5 km E Potter, 41°04.310'N, 103°17.787'W, metal tank 6.1 m in diameter in rolling hills, one 
bat captured: 9 September 2010—none captured (net removed before midnight); 30 September 2010, p.m.— M. 
ciliolabrum (one adult male [UNSM 30096]). 

(9) 16.2 km S, 1.5 km E Potter, 41°04.316'N, 103°17.780'W, adjacent to an abandoned rock house in rolling 
hills, no bats captured on nights of 9 September 2010 (net removed before midnight) and 30 September 2010, 
including after midnight. 

(10) 14.0 km S, 13.4 km E Potter, 41°05.469'N, 103°09.278'W, metal tank 7.6 m in diameter in rolling hills, no 
bats captured on night of 2 September 2010, including after midnight. 

(11) 6 km S, 13.4 km E Potter, 41°09.785'N, 103°09.300'W, metal tank 1.8 m in diameter in rolling hills, no bats 
captured on nights of 1 August and 1 September 2010, including after midnight. 

(11) 5.2 km S, 13.7 km E Potter, 41°10.342'N, 103°09.045'W, metal tank 2.4 m in diameter in rolling hills, no 
bats captured on night of 1 September 2010 (net removed before midnight). 

(11) 6 km S, 14 km E Potter, 41°09.796'N, 103°08.976'W, earthen tank 25 by 8 m in length in rolling hills, two 
bats captured: 1 August 2010, p.m.— M. ciliolabrum (two lactating females [UNSM 30095]). 

Lodgepole Creek in Kimball County 

(4) Intersection of Lodgepole Creek and Highway 71,41°14.889'N, 103°39.767'W, along creek, two bats captured: 
4 September 2010, p.m.— E.fuscus (one adult female [UNSM 30087]), L. noctivagans (one adult male [UNSM 
30077]), and Myotis ciliolabrum (observation only). 


200 


Clyde Jones Memorial Volume 


Lodgepole Creek in Cheyenne County 

(8) 1.3 km S, 4.5 km E Potter, Lodgepole Creek, 41°12.356'N, 103°15.654'W, along creek, two bats captured: 5 
October 2010, p.m.— L. noctivagans (two adult females [UNSM 30080]). 

(12) 2.9 kmS, 14.5 km E Potter, Lodgepole Creek, 41° 11.510'N, 103°08.473'W, along creek, 17 bats captured: 30 
June 2010, p.m.— E.fuscus (two pregnant females [UNSM 30085], two lactating females, two adult females), and 
M. ciliolabrum (one pregnant female [UNSM 30092]); 1 July 2010, a.m.— E.fuscus (two pregnant females, one 
adult female); 3 September 2010, p.m.— E.fuscus (one adult female, one young-of-the-year male), L. noctivagans 
(one adult male [UNSM 30076]), and Myotis ciliolabrum (observation only); 6 September 2010, p.m. —Myotis 
ciliolabrum (observation only); 7 September 2010, a.m.— L. noctivagans (one adult male, one adult female); 30 
September 2010, p.m.— M. ciliolabrum (one adult male); 1 October 2010, a.m.— L. noctivagans (one adult male). 

(13) 1.1 km S, 0.7 km E Lodgepole, Lodgepole Creek, 41°08.338'N, 102°37.724'W, along creek, two bats captured: 
1 October 2010, p.m.— L. noctivagans (one adult female, one adult male [UNSM 30082]). 

Specimens collected prior to our study 

(1) Kimball County, Gross-Wilkerson Ranch, 7.56 km S, 12.39 km W Bushnell Post Office, T14N, R59W, Sec. 
24 SW%, three specimens: 15 July 2004— M. ciliolabrum (one female [UNSM 28970]); 17 July 2004— M. cili¬ 
olabrum (one lactating female [UNSM 29175]); 4 August 2004— M. ciliolabrum (one female [UNSM 30296]). 

(3) Kimball County, T16N, R56W, Sec. 26 (center of section is 11.2 km north-northwest of town of Kimball), 
one specimen: 17 May 1988— L. noctivagans (one female [UNSM 19369]). 


(4) Kimball County, Kimball area, one specimen: 16 May 1964— M. ciliolabrum (one male [UWYMV 124]). 


Molecular Systematics and Phylogeography of Peromyscus nudipes 

(Cricetidae: Neotominae) 


Robert D. Bradley, Maria Nuhez-Tabares, Taylor J. Soniat, Sara Kerr, Russell W. Raymond, and 

Nicte Ordonez-Garza 


Abstract 

The taxonomic status of Peromyscus nudipes has been problematic, with morpho¬ 
metric studies placing P. nudipes in synonymy with P. mexicanus, whereas molecular 
studies have indicated that P. nudipes may be an independent evolutionary lineage. 
To address this conundrum, DNA sequences from the mitochondrial cytochrome-^ 
gene, representing all but one of the currently recognized members of the P. mexica¬ 
nus species group, were obtained from GenBank or generated as part of this study. In 
addition, samples representing closely related species were included as reference and 
comparative samples. DNA sequences were analyzed under a Bayesian Inference 
model to infer phylogenetic relationships. Further, genetic distances were estimated 
and used to determine levels of genetic divergence among taxa. Results indicated that 
samples formerly recognized as P. nudipes nudipes from south-central Costa Rica and 
northern Panama formed a monophyletic group, whereas samples formerly assigned 
to P. nudipes Hesperus and P. nudipes orientalis grouped with samples currently rec¬ 
ognized as P. nicaraguae. Levels of genetic divergence estimated from samples of P. 
nudipes nudipes indicated that it was among the most divergent taxa residing in the P. 
mexicanus species group. Further, differences in distribution and elevation depicted 
a separation of samples representing P. nudipes nudipes and P. mexicanus. Together 
these data suggest that P. nudipes is a valid species and that its distribution should be 
restricted to the high elevation, montane forests of the Cordillera de Talamanca located 
in south-central Costa Rica and northern Panama. 

Key words: cytochrome-b gene, Mesoamerica, molecular systematics, Peromys¬ 
cus, P. mexicanus species group, P. nudipes 


Introduction 


The systematic status of Peromyscus mexicanus 
(Rodentia, Cricetidae) and its allies comprising the P. 
mexicanus species group (Osgood 1909; Carleton 1989; 
Musser and Carleton 2005) has been problematic for 
several decades. This complex, distributed throughout 
southern Mexico and Central America, occupies a 
diversity of habitats and elevation ranges and presum¬ 
ably its evolutionary history has been influenced by 
numerous biogeographic and climatic events (Perez 
Consuegra and Vazquez-Dominguez 2015 and citations 
therein). The abiotic and biotic features of this region 
make it one of the most biologically diverse regions in 


the world and certainly these factors have impacted the 
diversification of Peromyscine rodents (Dawson 2005). 

Since Osgood (1909), numerous authors have 
revised and improved our understanding of the P. 
mexicanus species complex (Hooper and Musser 1964; 
Hooper 1968; Musser 1969; Huckaby 1980; Carleton 
1989; Rogers and Engstrom 1992; Ordonez-Garza et al. 
2010); however, incomplete sampling from the entire 
distribution or exclusion of critical taxa has hampered 
a global assessment of this species complex. Perez 
Consuegra and Vazquez-Dominguez (2015) based on 


202 


Clyde Jones Memorial Volume 


findings from analyses of mitochondrial cytochrome-/? 
gene (Cytb) sequences, provided the most recent taxo¬ 
nomic assessment of the P. mexicanus species group. 
This extensive study resulted in the proposed elevation 
of three junior synonyms to species status (P. nicara- 
guae, P. salvadorensis, and P. tropicalis ), resulting in an 
increase of the number of species in the group to 11 (P. 
grandis, P. guatemalensis, P. gymnotis, P. mexicanus, 
P. nicaraguae, P. nudipes, P. salvadorensis, P. stirtoni, 
P. tropicalis, P. yucatanicus, and P. zarhynchus ). 

Although the study by Perez Consuegra and 
Vazquez-Dominguez (2015) resulted in the broadest 
geographic coverage to date, their study included very 
few samples located south of Honduras. In particular, 
only two samples associated with P. nudipes were 
included in their study - one from San Jose, Costa 
Rica (GenBank accession number KP284425) and 
the specimen reported in Miller and Engstrom (2008) 
from Cartago, Costa Rica (GenBank accession number 
EF989992). Although P. nudipes does not have a broad 
geographic distribution (see Hall 1981), the geographic 
features of Costa Rica and Panama (Gutierrez-Garcia 
and Vazquez-Dominguez 2013) preclude an in-depth 
analysis of genetic variation within this species relative 
to its geographic distribution and to other P. mexicanus 
group members located throughout Mesoamerica. 

Allen (1891) described P. nudipes based on a 
single specimen collected from La Carpintera, Costa 
Rica. Osgood (1909) examined material from ad¬ 
ditional localities and the data supported Allen’s 
description but it was noted that P. nudipes appeared 
to be intermediate between P. guatemalensis and P. 
mexicanus. Several years later, Goodwin (1938) and 
Harris (1940) described two additional subspecies of 
P. nudipes from Costa Rica; P. n. orientalis and P. n. 
hesperus, respectively. Goodwin (1946) examined 
additional material from Costa Rica and Panama and 
postulated that the type specimen is not a “typical rep¬ 
resentative” of P. nudipes but that specimens collected 
from Volcan Irazu (~ 2,850 m) were more reflective 
examples of the species. In Goodwin’s (1946) synopsis, 
P. n. orientalis and P. n. hesperus were determined to 
be smaller in size and appeared to occupy lower eleva¬ 
tion habitats than did P. n. nudipes. Further, Goodwin 
(1946) surmised that P. n. orientalis and P. n. hesperus 
might be aligned with P. mexicanus saxatilis ; leaving 
only P. n. nudipes as a representative of the species. 


Hooper (1968) continued to treat P. nudipes as a spe¬ 
cies although he echoed Osgood’s (1909) position that 
it could fit into either P. guatemalensis or P. mexicanus. 
Huckaby’s (1980) detailed revision of the P mexicanus 
species group resulted in P. nudipes being synonomized 
with P. mexicanus. Although Hall (1981) recognized 
P. nudipes (and the subspecific divisions), it is likely 
that he was either unaware of or ignored Huckaby’s 
revisionary study. More recent synopses (Carleton 
1989; Musser and Carleton 2005; Trujano-Alvarez 
and Alvarez-Castaneda 2010) followed Huckaby’s 
recommendation of placing P. nudipes in synonomy 
with P. mexicanus. 

Although the morphologic data argue for syn- 
onymizing all of P. nudipes {hesperus, orientalis, and 
nudipes), or at least P. n. hesperus and P. n. orientalis, 
with P. mexicanus', the genetic data suggest that dif¬ 
ferentiation exists between the two taxa. For example, 
despite extreme conservation among autosomes, P. 
nudipes differs from P. mexicanus (and some other P. 
mexicanus group members) in the morphology of the X 
and Y-chromosomes (Hsu andArrighi 1968; Rogers et 
al. 1984; Smith et al. 1986). Further, in the allozymic 
study of Rogers and Engstrom (1992), P. nudipes, P. 
mexicanus, P. guatemalensis, P. yucatanicus, and P. 
zarhynchus were found to be genetically similar but 
differed based on the presence of autapomorphic al¬ 
leles. Analyses of mitochondrial DNA sequence data 
(Miller and Engstrom 2008, Perez Consuegra and 
Vazquez-Dominguez 2015) demonstrated that samples 
historically assignable to P. n. nudipes (La Trinidad de 
Dota, and Cerro de la Muerte, Costa Rica) were strongly 
differentiated from samples of P. mexicanus. 

The goals of this study were to: 1) examine 
genetic variation in the Cytb gene in samples of P. 
nudipes and P. nicaraguae, including specimens for¬ 
merly assigned to P. n. hesperus and P. n. orientalis, 
from Costa Rica, Honduras, Nicaragua, and Panama; 
and 2) attempt to resolve the taxonomic status of P. 
nudipes relative to other members of the P. mexicanus 
species group. DNA sequences generated herein were 
combined with sequences from Perez Consuegra and 
Vazquez-Dominguez (2015) and sequences obtained 
from GenBank to better assess the genetic variation 
among members of the P. mexicanus group from the 
more southern portion of its range. 


Bradley et al.—Systematics of Peromyscus nudipes 


203 


Materials and Methods 


Samples .—Tissue samples were obtained from 
51 specimens collected from naturally-occurring 
populations in Honduras, Mexico, and Nicaragua or 
borrowed from museum collections (Localities 1-18, 
Fig. 1; Appendix). DNA sequences from an additional 
173 individuals were included as internal references or 
used for outgroup comparisons based on the studies 
of Bradley et al. (2007), Miller and Engstrom (2008), 
Ordonez-Garza et al. (2010), and Perez Consuegra 
and Vazquez-Dominguez (2015). Specimens were 
collected following methods outlined in the ASM 
Guidelines (Sikes et al. 2016) and approved by the 


Texas Tech University Animal Care and Use Commit¬ 
tee. Specimen numbers and collection localities are 
listed in the Appendix. 

Sequence data .—Methods for obtaining DNA 
sequencing data follow that of Bradley et al. (2007, 
2015) with minor modifications; as summarized below. 
Mitochondrial DNA was isolated from approximately 
0.1 g of tissue using a DNeasy kit (Qiagen, Valencia, 
California). The entire Cytb gene (1,143 bp) was ampli¬ 
fied using the polymerase chain reaction method (PCR, 
Saiki et al. 1988) and the following primers: MVZ05 


f**. 11 


SWr'W 


i5<w"s 


P. nicaraguae 


P. nudipes orientalis 


P. nudipes hesperus 


200 km 


P. nudipes nudipes 


90°Q’ir\Y 


85 c ’0'fl”W 


Figure 1. Distribution of selected populations and species of the Peromyscus mexicanus species group from 
Mesoamerica. Closed circles represent sampling localities and numbers refer to samples listed in the Appendix. 
Open diamond, star, and triangle represent the approximate location of the type localities for P. nudipes nudipes, P. 
nudipes hesperus, and P. nudipes orientalis , respectively. 


204 


Clyde Jones Memorial Volume 


(Smith and Patton 1993) and PER03 ’ (Tiemann-Boege 
et al. 2000). Thermal profiles for PCR were as follows: 
initial denaturation at 95°C for 2 min, followed by 35 
cycles of denaturation at 95°C for 1 min, annealing 
at 51°C for 1 min, and extension at 72°C for 2 min, 
with a final extension at 72°C for 7 min. Most PCR 
products were purified with ExoSAP-IT (Asymetrix, 
Santa Clara, California). Primers used to cycle se¬ 
quence the products included: WDRAT1100, 400R, 
700H, and NEO700L (Peppers and Bradley 2000) and 
400F, (Tiemann-Boege et al. 2000). Cycle sequencing 
reactions were purified using isopropanol cleanup 
protocols and were analyzed with an ABI 3100-Avant 
automated sequencer and ABI Prism Big Dye version 
3.1 terminator technology (Applied Biosystems, Fos¬ 
ter City, California). Resulting sequences (of various 
lengths) were aligned and proofed using Sequencher 
4.0 software (Gene Codes, Ann Arbor, Michigan); chro¬ 
matograms were examined to verify all base changes. 
Cytb sequences obtained in this study were deposited 
in GenBank and are listed in the Appendix. 

Using the phylogenetic relationships of the genus 
Peromyscus proposed by Bradley et al. (2007) and Platt 
et al. (2015), Reithrodontomysfulvescens was selected 
as the outgroup taxon for sequence analyses. Ten of 
the 11 species putatively assigned to the P. mexicanus 
species group (Bradley et al. 2007; Perez Consuegra 
and Vazquez-Dominguez 2015 — P. grandis, P. gua- 
temalensis, P. gymnotis, P. mexicanus, P. nicaraguae, 
P. nudipes, P. salvadorensis, P. stirtoni, P. tropicalis, 
and P. zarhynchus ) were included in the analyses; 
samples were not available for P. yucatanicus. In ad¬ 
dition, samples of P. furvus, P. mayensis, P. megalops, 
P. melanocarpus, P melanophrys, and P. perfulvous 
were included based on their potential affiliations to 
the P. mexicanus species group. Further, samples of P. 
boylii, P. leucopus, and P. ochraventer were included 


as internal standards. All analyses were performed in 
PAUP* (Version 4.0al49; Swofford 2002). 

Eighty-eight maximum likelihood models were 
evaluated using MODEFTEST (Darriba et al. 2012) 
in order to determine the model of DNA evolution 
best fitting the data. The Akaike information criterion 
identified HKY+I+G as being the most appropriate 
model, relative to complexity of model, for this dataset. 
This model generated significantly better likelihood 
scores (-InF = 13002.3955) than all other models and 
included the following parameters: base frequencies 
(A= 0.3525, C = 0.3080, G = 0.0838, and T = 0.2557); 
proportion of invariable sites (I = 0.4590); and gamma 
distribution (G = 0.8810). 

Bayesian inference methods (MrBayes; Huelsen- 
beck and Ronquist 2001) were used under a maximum 
likelihood framework and the HKY+I+G model of 
evolution. Model parameters were estimated within 
the analysis. The analysis was run with the following 
options: 4 Markov-chains, 10 million generations, and 
sample frequency = every 1,000 th generation. Fol¬ 
lowing a visual inspection of likelihood scores, the 1 st 
1,000 trees were discarded and a consensus tree (50% 
majority rule) was constructed from the remaining 
trees. Clade probabilities values, indicative of nodal 
support, were estimated and only values >0.95 were 
considered as evidence for statistical support. 

The Kimura 2-parameter model of evolution 
(Kimura 1980) was used to calculate genetic distances 
between selected taxa. These genetic distances were 
used: 1) as a direct comparison to values reported in 
Perez Consuegra and Vazquez-Dominguez (2015) and 
2) to assess levels of genetic divergence following cri¬ 
teria outlined in Bradley and Baker (2001) and Baker 
and Bradley (2006). 


Results 


A total of 226 DNA sequences (of various 
lengths) representing the P. mexicanus species group, 
other closely related species groups, and the outgroup 
taxon were analyzed using Bayesian inference methods. 
In this analysis, only clade probability values (CPV) 
>0.95 were considered as evidence for nodal support. 


The resulting topology (Fig. 2) depicted a large clade 
(Clade A, CPV = 1.00) that included the following taxa: 
P. mayensis, P. megalops, P. stirtoni, P. melanocarpus, 
P. melanophrys, P. perfulvous and 9 species (Clade B, 
CPV = 1.00) referred, herein, as the mexicanus species 
group ( sensu stricto). Given that the phylogenetic 


Bradley et al.—Systematics of Peromyscus nudipes 


205 


Reithrodontomys fulvescens 


A 


0.97 

Nl 


Peromyscus ochraventer (n = 2) 

- Peromyscus boylii 

Peromyscus megalops 

- Peromyscus stirtoni (n = 1) 

Peromyscus grandis (n = 1) 

— Peromyscus grandis (n = 9) 
Peromyscus guatemalensis (n = 7) 
Peromyscus guatemalensis (n = 12) 


B 


Peromyscus guatemalensis (n = 36) 

* Peromyscus salvadorensis (n = 19) 

* Peromyscus salvadorensis (n = 13) 
Peromyscus guatemalensis (n = 4) 
Peromyscus nicaraguae (Loc 5, n = 4) 


C 


'i' 

iL 


Peromyscus nicaraguae (Loc 4, 6, and 8, n = 4) 
Peromyscus nicaraguae (Loc 6, n = 1) 
Peromyscus nicaraguae (Locs 7 and 9, n = 15) 

- Peromyscus nicaraguae (Loc 15) 


0.95 


- Peromyscus nicaraguae (Locs 1—5, n = 18) 

' i-—- Peromyscus nicaraguae (Loc 13, n = 2) 

—-> 4 . p ( > 


•f 

rC 


Peromyscus nicaraguae (Locs 10--14, n = 7) 

Peromyscus zarhynchus (n = 11) 

— Peromyscus zarhynchus (n = 6) 

- Peromyscus zarhynchus (n = 9) 


Peromyscus gymnotis (n = 7) 


0.98 


Peromyscus mexicanus (n = 7) 


Peromyscus mexicanus (n = 6) 


D 


Peromyscus nudipes (Loc 17, n = 1) 


Peromyscus nudipes (Loc 18, n = 2) 
Peromyscus nudipes (Loc 16, n = 1) 

- Peromyscus tropicalis (n = 2) 


- Peromyscus melanocarpus (n = 1) 

.j. - Peromyscus melanophrys (n = 2) 

- Peromyscus perfulvous (n = 1) 

--- Peromyscus mayensis (n = 4) 

I- Peromyscus leucopus (n = 1) - 0 .03 Substitutions/Site 

- - - Peromyscus jurvus (n = 4) 

Figure 2. Phylogenetic tree generated using Bayesian methods (MrBayes; Huelsenbeck and Ronquist 2001) and the 
HKY+I+G model of evolution. Clade probability values are shown above branches values (values of 1.00 are indicat¬ 
ed by an asterisk). Terminal taxa represented in bold font are indicative of samples formerly assigned to Peromyscus 
nudipes nudipes , P. nudipes hesperus, and P. nudipes orientalis. Localities (Loc) and samples sizes (n) are provided 
in parentheses and correspond to the specimens examined in the Appendix. 


206 


Clyde Jones Memorial Volume 


status of P. mayensis and P. stirtoni remain unclear 
and that P. megalops, P. melanocarpus, P. melanoph- 
rys, and P. perfulvus have been shown to be affiliated 
with other species groups (Musser and Carleton 2005; 
Bradley et al. 2007), a detailed presentation of these 
results will be restricted to the membership of clade B. 

Within Clade B, individuals formed monophy- 
letic clades within the boundaries of their respective, 
designated species except for samples of P. guate- 
malensis and P. mexicanus. Samples of P. n. hesperus 
(Localities 12 and 13), P. n. orientalis (Locality 14) 
and P. n. nudipes (Locality 15) were members of a 
clade containing individuals of P. nicaraguae (Clade 

C, Localities 1-11) from Honduras and Nicaragua; 
whereas the remaining samples of P. n. nudipes (Clade 

D, Localities 16-18) formed an independent clade near 
the base of Clade B. 

Genetic divergence values (Table 1) were esti¬ 
mated for representative individuals using the Kimura 
2-parameter model of evolution (Kimura 1980). In this 


analysis, only a single representative of each population 
was included in the analyses to prevent an underestima¬ 
tion of genetic differentiation. Further, based on the 
results of the Bayesian analysis, only samples from 
Localities 16-18 were included as representatives ofP 
nudipes, samples from localities 10-15 were treated as 
samples of P. nicaraguae. Average genetic distances 
among samples, representing each species ranged from 
1.28% ( P. gymnotis ) to 4.17% (P zarhynchus ); samples 
of P. tropicalis differed by 0.09%, however, only 2 
individuals were available for analysis and they were 
from the same population. Genetic divergence values 
between species ranged from 4.65% ( P. guatemalensis 
and P. salvadorensis ) to 11.13% (P gymnotis and P. 
nudipes). Pairwise comparison to other species in the 
P. mexicanus group revealed that P. nudipes differed 
by values ranging from 9.20% (P salvadorensis) 
to 11.13% (P gymnotis). Overall, levels of genetic 
divergence between P nudipes and others species in 
the P mexicanus were similar to or exceeded values 
resulting from pairwise comparisons of all P mexicanus 
groups members. 


Discussion 


A monophyletic clade (Clade B, Fig. 2) contain¬ 
ing members of the P mexicanus group was obtained 
from the Bayesian Inference analysis. A sensu stricto 
interpretation of this group indicates that the P. mexi¬ 
canus group should include the following species: P. 
grandis, P. guatemalensis, P. gymnotis, P. mexica¬ 
nus, P. nicaraguae, P. nudipes, P. salvadorensis, P. 
tropicalis, and P. zarhynchus. Although samples of P. 
yucantanicus were not available for inclusion in this 
study, several lines of evidence (Huckaby 1980; Rogers 
et al. 1984; Smith et al. 1986; Rogers and Engstrom 
1992) support its inclusion into the P mexicanus species 
group. Phylogenetic relationships between most spe¬ 
cies were resolved, however, P. tropicalis, P. nudipes, 
and a clade containing samples of P. gymnotis and P. 
mexicanus could not be placed within the context of 
the remaining species. Although beyond the scope of 
this study, it is apparent that multiple species probably 
reside in what is now recognized as P guatemalensis 
and P mexicanus. Further, in agreement with Carleton 
(1989) and Bradley et al. (2007), it appears that the 
phylogenetic relationships obtained from this study 


indicate that P mayensis and P melanocarpus are prob¬ 
ably not members of the P mexicanus species group. 

Based on the results of the Bayesian Inference 
analysis and magnitude of genetic divergence re¬ 
ported herein, P nudipes is a valid species; however, 
we suggest a more restricted view of this taxon than 
previously described. In agreement with previous au¬ 
thors (Goodwin 1946; Hooper 1968; Huckaby 1980; 
Carleton 1989; Musser and Carleton 2005; Trujano- 
Alvarez and Alvarez-Castaneda 2010), samples of P 
nudipes orientalis and P nudipes hesperus are more 
closely related to P guatemalensis or P mexicanus. In 
the phylogenetic analysis, presented herein, samples 
from northern Costa Rica (Localities 12-14) formerly 
considered to be representative of P nudipes orienta¬ 
lis and P nudipes hesperus were included in a large 
clade (Clade C) containing samples from Honduras 
and Nicaragua (Localities 1-11). The samples from 
Localities 1-11 historically have been recognized as 
P m. saxatilis (formal synonymy provided by Trujano- 
Alvarez and Alvarez- Castaneda 2010); portions of 


Bradley et al.—Systematics of Peromyscus nudipes 


207 


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this taxon recently were elevated to P. nicaraguae by 
Perez Consuegra and Vazquez-Dominguez (2015). In 
addition, the sample from Locality 15, formerly aligned 
with P. nudipes nudipes, also was embedded within the 
clade containing representatives of P. nicaraguae and 
should be recognized as such. Samples from Localities 
16-18 (south-central Costa Rica and Panama) formed 
a monophyletic clade (clade D) positioned near the 
base of the P. mexicanus group topology and distantly 
removed from members of clade C. The samples 
comprising clade C historically were considered to 
be examples of P. nudipes nudipes and should retain 
that name based on phylogenetic principles and levels 
of genetic divergence from other members of the P. 
mexicanus species group. 

In resolving this conundrum, besides the genetic 
differentiation reported herein, it appears that elevation 
may be a key factor in further distinguishing samples of 
P. nudipes from those now included in P. nicaraguae. 
The type locality for P. nudipes nudipes (La Carpin- 
tera, Costa Rica) was reported by J. A. Allen (1891) 
to be approximately 1,818 m in elevation. Elevations 
reported for the type localities for P. nudipes Hesperus 
(Hacienda Santa Maria, Costa Rica, about 15 mi NE 
of Liberia; Harris 1940) and P. nudipes orientalis (El 
Sauce Peralta, Costa Rica; Goodwin 1938) are ~ 970 
m and 303 m, respectively. Elevations at the type 
localities for both P nudipes Hesperus and P nudipes 
orientalis appear to be more similar to the low to mid 
elevation ranges typically associated with P. mexicanus 
saxitalis (now considered to be P. nicaraguae ). Con¬ 
versely, the samples of P nudipes nudipes, examined 
herein, were from much higher elevations (2,500 m 
- Locality 16, 2,325 m - Locality 17, and 2,000 m - 
Locality 18). It appears that P. nudipes is restricted 
to the montane pine-oak forests of the Cordillera de 
Talamanca of southern Costa Rica and northwestern 
Panama, whereas P. nicaraguae (formerly P. nudipes 
Hesperus and P. nudipes orientalis ) occupies the mid 
to low elevation tropical forests of the Central, Tilaran, 
and Guanacaste Mountains in the north-central regions 
of Costa Rica, northward to Nicaragua and Honduras. 
This hypothesis needs to be tested by obtaining more 
samples from trapping to determine if taxa are restricted 
to these ecosystems. 

The difference in geological age of these two 
ranges may have been a contributing factor to the 


208 


Clyde Jones Memorial Volume 


differentiation of P. mexicanus and P. nudipes. The 
younger and more northern ranges (Central, Tilaran, 
and Guanacaste) were formed from volcanic activity 
during the Quaternary (Soto and Alvarado 2006; Carr 
et al. 2007; Gutierrez-Garcia and Vazquez-Dominguez 
2013). Conversely, the older and more southern range 
(Cordillera de Talamanca) rapidly uplifted during the 
late Pliocene to early Pleistocene and had little volcanic 
activity during the Quaternary (Johnston and Thorkel- 
son 1997). The forests of the Cordillera de Talamanca 
are considered the largest continuous forested area in 
Costa Rica (Tobler 2002) and are characterized by 
steep slopes and numerous small inter-montane valleys 
(Gonzalez-Maya et al. 2009). This habitat (classified 
as Tropical Montane Cloud Forests, Kappelle 1996) 
is dominated by two species of oak (Tobler 2002) 
and typically ranges from 2,000 to 3,200 m. Ad¬ 
ditional sampling is needed from the high elevation 
areas (>2,000 m) in central and southern Costa Rica, 
as well as northern Panama, to resolve the elevation 
and habitat preferences and differences between P. 
mexicanus and P. nudipes. Further, special attention 
is needed for samples from the southern Central Range 
near Volcan Irazu, which Goodwin (1946) ascertained 
as being more closely aligned with P. nudipes. It may be 
that P. nudipes occurs at high elevations in this region. 

The decision to resurrect the name P. nudipes (J. 
A. Allen 1891) from synonymy with P. mexicanus (see 
Trujano-Alvarez and Alvarez-Castaneda 2010) was 
based on the following. First, a comprehensive phylo¬ 
genetic analysis that included all currently recognized 
members of the P. mexicanus group, representative of 
presumed closely related species, as well as, a sampling 
scheme that included samples from throughout the en¬ 
tire range of the P. mexicanus species group provided 
data that supported P. nudipes as a monophyletic as¬ 
semblage. This monophyletic group contained samples 


from high elevation locations in south-central Costa 
Rica and Panama that resembled the initial supposition 
and description of P. nudipes by Allen (1891). Once 
these high elevation samples (Localities 16-18) were 
included with the broad geographic coverage reported 
in the taxonomic revision by Perez Consuegra and 
Vazquez-Dominguez (2015), it became apparent that 
P. nicaraguae and P. nudipes represented independent 
evolutionary complexes. 

Second, examination of the genetic distance 
dataset indicated that P. nudipes was among the most 
genetically divergent members of the P. mexicanus 
species group. Samples of P. nudipes differed from 
other P. mexicanus group members by average genetic 
distances ranging from 9.26% to 11.13%. As a com¬ 
parison, the sister species P. melanophrys and P. perful- 
vus differed by 7.92% (this study). In addition, these 
values exceeded those estimated from other groups of 
closely related Peromyscus species (Durish et al. 2004; 
Bradley et al. 2004,2014) indicating either a more rapid 
rate of molecular divergence or a longer elapsed time 
period since sharing a most recent common ancestor. 
Interpretation of the phylogenetic and genetic distance 
data under the auspices of the Genetic Species Concept 
(see Bradley and Baker 2001 and Baker and Bradley 
2006) argues that P. nudipes represents a genetic species 
within the P. mexicanus species group. 

In conclusion, it is suggested that 1) P. nudipes, 
restricted to samples from south-central Costa Rica 
and northern Panama, be reinstated as a species, 2) 
samples of P. nudipes Hesperus and P. nudipes orien- 
talis should be subsumed into P. nicaraguae, and 3) 
further sampling be conducted in the montane regions 
of central Costa Rica to better diagnose the distribution 
and habitat differences associated with P. nudipes and 
P. nicaraguae. 


Acknowledgments 


We thank E. K. Roberts, J. Q. Francis, and 
L. L. Lindsey for reviewing earlier versions of this 
manuscript. We appreciate the generous help from 
J. O. Matson, S. Parsons, R. and M. Eckerlin, W. and 
J. Bulmer for assistance and supporting fieldwork in 
Guatemala. We thank the Field Methods classes of 


2001 and 2004 for assistance in obtaining specimens 
from Honduras. Franklin Herrera at Consejo Na- 
cional de Areas Protegidas (CONAP) of Guatemala 
and Enrique Fajardo at the Ministerio de Ambiente y 
Recursos Naturales of El Salvador provided permits 
and valuable assistance during field season. Tissue 


Bradley et al.—Systematics of Peromyscus nudipes 


209 


samples were provided by the Natural Science Research 
Laboratory, Museum of Texas Tech University, Natural 
History Museum, University of Kansas, and Zadock- 
Thompson Natural History Collection, University of 


Vermont. Partial support for this research was provided 
by two NIH grants (DHHS A14143 5-01 to RDB; 2 S06 
GM55337-05 to SK). 


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Bradley et al.—Systematics of Peromyscus nudipes 


211 


Addresses of authors: 


Robert D. Bradley 

Sara Kerr 

Department of Biological Sciences and Museum 
Texas Tech University 

Lubbock, TX 79409-3131 
robert. bradley @ ttu. edu 

Department of Biology 

University of the Incarnate Word 
4301 Broadway 

San Antonio, TX 78209 

Maria Nunez-Tabares 

Russell W. Raymond 

Department of Biological Sciences 

Texas Tech University 

Lubbock, TX 79409-3131 

Department of Biology 

University of the Incarnate Word 
4301 Broadway 

San Antonio, TX 78209 

Taylor J. Soniat 

Nicte Ordonez-Garza 

Department of Biological Sciences 

Texas Tech University 

Lubbock, TX 79409-3131 

Department of Biological Sciences 
Texas Tech University 

Lubbock, TX 79409-3131 


212 


Clyde Jones Memorial Volume 


Appendix 

Specimens examined in the DNA sequencing portion of this study. Taxonomic designations following and the 
results of Bradley et al. (2007), Perez Consuegra and Vazquez-Dominguez (2015), and those presented herein. 
For each specimen reported herein the collection locality, museum catalogue number (to left of slash) and Gen- 
Bank accession number (to right of slash) are provided in parentheses. Abbreviations for museum acronyms 
follow Hafner et al. 1997). For most sequences obtained from GenBank only the accession number is provided. 
Abbreviations are as follows: Museum of Natural History (KU), Museum of Southwestern Biology (MSB), 
Museum of Texas Tech University (TTU), United States National Museum of Natural History (USNM), and 
Zadock Thompson Natural History Collection, University of Vermont (ZTNHC). If museum catalog numbers 
were unavailable, specimens were referenced with the corresponding TK number (special number of the Museum 
of Texas Tech University). Localities corresponding to Figure 1 are provided in parentheses. 

Peromyscus mexicanus species group: 

Peromyscus grandis .—GUATEMALA: Alta Verapaz; Chelemha Yalijux Mountain (USNM569843/KX998940, 
USNM569910/KX998941). Sequences from GenBank: GQ461919, GQ461920, GQ461921, GQ461922, 
GQ461923, GQ461924, GQ461925, KP284306. 

Peromyscus guatemalensis .—GUATEMALA: Huehuetenango; 9 km NNE Cuilco, El Retiro (USNM570208/ 
KX998915, USNM570213/KX998916, USNM570220/KX998917),10 Km. NNE Cuilco, El Retiro (USNM570233/ 
KX998918, USNM570234/ KX998919, USNM570235/KX998920, USNM570236/KX998921, USNM570237/ 
KX998922, USNM570239/KX998923, USNM570240/KX998924, USNM570241/KX998925, USNM570242/ 
KX998926, USNM570243/KX998927). Sequences from GenBank: EF281171, EF281172, GQ461926, 
GQ461927, GQ461928, GQ461929, GQ461930, GQ461931, GQ461932, GQ461933, GQ461934, GQ461935, 
KP284351, KP284352, KP284353, KP284354, KP284355, KP284356, KP284357, KP284358, KP284359, 
KP284360, KP284361, KP284362, KP284363, KP284364, KP284365, KP284366, KP284367, KP284368, 
KP284369, KP284370, KP284371, KP284372, KP284373, KP284374, KP284375, KP284376, KP284377, 
KP284377, KP284378, KP284379, KP284380, KP284381, KP284382, KP284383, KP284384. 

Peromyscus gymnotis .—Sequences from GenBank: EF028169, EF028170, KP284385, KP284386, KP284387, 
KP284388, KP284389. 

Peromyscus mexicanus .—EL SALVADOR: Santa Ana; Parque Nacional Montecristo (TTU61014/KX998944. 
GUATEMALA: Chiquimula, 8 Km SW Esquipulas, Plan de la Arada, (TTU125805/KX998942, TTU125806/ 
KX998943). MEXICO: Chiapas, 25 km S, 3 km N Ocozocoautla (TTU104622/KX998938); Veracruz; 6.7 km 
NE, 13.5 km SE Perote (TTU105005/KX998945). Sequences from GenBank: EF028174, HQ269736, KP284422, 
KP284423, KP284424, KJ526415, AY376425, JX910118. 

Peromyscus nicaraguae .—COSTARICA: Cartago; Capellades; (Loc 15, EF989991); Guanacaste; Volcan Santa 
Maria (Loc 13, EF989993); Heredia: 2 kmNE Getzemani (Loc 14, AY041200); Puntarenas; Monte Verde Biologi¬ 
cal Station (Loc 13, KU142102/KX998930, KU143321/KX998931, EF989994). HONDURAS: Colon: Trujillo; 
Parque Nacional Capiro y Calentura (Loc 1, TTU104186/KX998935); Comayagua; Parque Nacional Cerro Azul 
Meambar (Loc 2, TTU104357/KX998934); Francisco Morazan; Parque Nacional La Tigra (Loc 5, TTU83698/ 
KX998948, TTU83731/KX998932, TTU83708/KX998963, KP284309, KP284310, KP284311, KP284312,); Olan- 
cho; Parque Nacional La Muralla (Loc 3, KP284314, KP284315, KP284316, KP284317, KP284318, KP284319, 
KP284320); Parque Nacional Sierra de Agalta, Babilionia Mountain (Loc 4, KP284321, KP284322, KP284323, 
KP284324, KP284325, KP284326, KP284327). NICARAGUA: Chinandega; Chichigalpa, Bella Vista (Loc 10, 


Bradley et al.—Systematics of Peromyscus nudipes 


213 


TTU105099/KX998959, TTU105096/KX998962); San Cristobal (Loc 11, TTU105108/KX998960, TTU105111/ 
KX998961); Madriz; San Lucas, Los Mangos (Loc 6, TTU9672/FJ214687); Jinotega; El Cua, Galope (Loc 7, 
TTU119600/KX998936, TTU119601/KX998937); Matagalpa; Selva Negra (Loc 9, TTU88195/KX998951, 
TTU93013/KX998954, TTU97009/KX998953, TTU97020/KX998947, TTU96980/KX998964, TTU105180/ 
KX998939, TTU105193/KX998955, TTU105190/KX998956, TTU105164/KX998958, TTU101386/KX998957, 
TTU101406/KX998933, TTU101409/KX998949, TTU101421/KX998950, TTU101441/KX998952), Posada 
Tisev (Loc 8,KP284313). 


Peromyscus nudipes .—COSTARICA: Cartago; Cerro de la Muerte, San Gerardo del Dota (Loc 17, EF989992); 
San Jose; 2.2 km E (by road) La Trinidad de Dota (Loc 16, KP284425). PANAMA: Chiriqui; Bugaba, La Amis- 
tad International Park, Las Nubes Ranger Station (Loc 18, MSB262229/KX998928, MSM262207/KX998929). 


Peromyscus salvadorensis .—Sequences from GenBank: KP284390, KP284391, KP284392, KP284393, KP284394, 
KP284395, KP284396, KP284397, KP284398, KP284399, KP284400, KP284401, KP284402, KP284403, 
KP284404, KP284405, KP284406, KP284407, KP284408, KP284409, KP284410, KP284411, KP284412, 
KP284413, KP284414, KP284415, KP284416, KP284417, KP284418, KP284419, KP284420, KP284421. 


Peromyscus stirtoni .—DQ973108. 

Peromyscus tropicalis .—Sequences from GenBank: KP284307, KP284308. 


Peromyscus zarhynchus .—GUATEMALA: Huehuetenango; Nenton 1 km NE (by road) of Yalmbojoch 
(USNM570460/KX998946). MEXICO: Chiapas; Yalentay UTM 15-524171-1852486 (TK93297/AY195800). 
Sequences from GenBank: EF028167, KP284328, KP284329, KP284330, KP284331, KP284332, KP284333, 
KP284334, KP284335, KP284336, KP284337, KP284338, KP284339, KP284340, KP284341, KP284342, 
KP284343, KP284344, KP284345, KP284346, KP284347, KP284348, KP284349, KP284350. 


Peromyscus mayensis .—Sequences from GenBank: DQ836300, DQ836301, EF989987, EF989988. 
Peromyscus melanocarpus .—Sequences from GenBank: EF028173. 

Outgroup and internal reference specimens: 

Peromyscus /wrvws.—AF270993, AF270995, AF271006, AF271026. 

Peromyscus megalops .—DQ000475. 

Peromyscus melanophrys .—AY322510, AY376424. 

Peromyscus perfulvus .—DQ000474. 

Peromyscus boylii .—DQ000478. 

Peromyscus leucopus .—DQ000483. 

Peromyscus ochraventer .—FJ214689, JX910119. 


Reithrodontomys fulvescens .—AF176257. 


An Inventory of Bats in Arch Canyon, San Juan County, Utah 

Tony R. Mollhagen and Michael A. Bogan 

Abstract 

Arch Canyon is located in southeastern San Juan County, Utah, southwest of 
Blanding, and west of Comb Ridge. During the summer of 2007, we conducted an 
inventory for bats in the canyon at the request of the Bureau of Land Management, 
which oversees management of the area. We were especially interested in the status of 
state species of concern, which include Fringed Myotis, Myotis thysanodes ; Allens Big- 
eared Bat, Idionycteris phyllotis', Townsends Big-eared bat, Corynorhinus townsendii; 
Spotted Bat, Euderma maculatum ; and Big Free-tailed Bat, Nyctinomops macrotis. 
We netted at 10 sites in the canyon and captured a total of 295 individual bats of 15 
species. The eight most common species of bats were Western Pipistrelle, Parastrellus 
hesperus (32.9% of total captures), Big Free-tailed Bat (23.4%), Brazilian Free-tailed 
Bat, Tadarida brasiliensis (12.5%), Big Brown Bat, Eptesicus fuscus (7.1%), Pallid 
Bat, Antrozouspallidus (5.4%), Yuma Myotis, Myotis yumanensis (4.7%), Spotted Bat, 
Euderma maculatum (3.7%), and California Myotis, Myotis californicus (3.4%). We 
captured 11 individual Euderma maculatum , a number that seems remarkable given 
the level of effort. The capture of 69 individual Big Free-tailed Bats, representing over 
23 percent of the total sample of Arch Canyon, is unusually high in comparison with 
other surveys in nearby areas. 

Key words: Arch Canyon, bats, Chiroptera, inventory, Utah 


Introduction 


The Colorado Plateau has attracted the attention 
of many scientists since J. W. Powell explored and 
mapped the canyon country of the Colorado River in 
1869 (Powell [reprinted] 1961). C. H. Merriam, V. 
Bailey, M. Cary, W. H. Osgood, and other employees of 
the Bureau of Biological Survey conducted biological 
explorations of the area in the late 1800’s. More recent 
studies of mammals found on the Colorado Plateau 
have included those by Durrant (1952) for Utah, Arm¬ 
strong (1972) and Fitzgerald et al. (1994), Armstrong et 
al. (2011) for Colorado, Findley et al. (1975) for New 
Mexico, and Floffmeister (1986) for Arizona. All of 
the aforementioned researchers and their work have 
contributed to our understanding of the fauna of the 
Colorado Plateau. 

Nonetheless, details of distribution and abun¬ 
dance for many species of plants and animals of the 


southeastern Utah canyon country are poorly known, 
and future management plans for such areas are depen¬ 
dent upon the availability of current information on the 
status of these species. It was in this context that the 
Monticello office of the Bureau of Land Management 
(BLM) requested specific information for selected 
species of small mammals occurring in canyons in 
southeastern Utah. BLM was specifically interested in 
acquiring information concerning bats in Arch Canyon. 

Arch Canyon is located in southeastern San Juan 
County, southwest of Blanding, and west of Comb 
Ridge. It is accessed from US Hwy 95, initially through 
tribal lands. There is an 8.9 mi (19.5 km), primitive, 
two-track trail from the canyon entrance to the National 
Forest Boundary. Motorized traffic entering the canyon 
must exit by the same route. Under normal circum¬ 
stances, several of the stream crossings are impassable 


215 


216 


Clyde Jones Memorial Volume 


for any except high-clearance, four-wheel-drive (4WD) 
vehicles. However, the canyon periodically is subject 
to flash flooding, thus rendering much of it inaccessible 
except by hiking. 

Arch Canyon appeals to a variety of interests. 
There are the eponymous arches in the upper canyon. 
There are a number of sites suitable for primitive camp¬ 
ing. Hiking is not difficult if walkers remain on the 
trail. Similarly, experienced drivers utilizing all-terrain 
(ATVs) or larger 4WD vehicles will find the trail some¬ 
times challenging, but usually not dangerous. There are 
the remains of at least three Anasazi dwellings along the 
canyon walls. Also, a resident fish species of concern, 
the Flannelmouth Sucker ( Catostomus latipinnis ) can 
be found in pools of the lower canyon. It is these last 
two attractions, plus the at least seasonally-resident 
chiropteran species, that raise concerns related to un¬ 
limited human access to the canyon. 

The bat species of concern include the Fringed 
Myotis, Myotis thysanodes; Allen’s Big-eared Bat, Idi- 
onycterisphyllotis; Townsend’s Big-eared Bat, Coryno- 
rhinus townsendii; Spotted Bat, Euderma maculatum; 
and Big Free-tailed Bat, Nyctinomops macrotis. All of 
these species occur on, but not necessarily throughout, 
the Colorado Plateau (e.g., Mollhagen and Bogan 1997, 
Bogan et al. 2006, O’Shea et al. 2011), and there is a 
high probability that these bat species are found in Arch 
Canyon. The possible impact of foot and vehicle traf¬ 
fic in association with recreational activities on these 
species is unknown. 

Durrant’s Mammals of Utah (1952), the first 
comprehensive report on mammals of Utah, provided 
statewide records for 17 species (1 erroneously) of bats. 
However, Durrant had little information on bats found 


in southeastern Utah. Armstrong (1974,1982) reported 
ten species of bats from Canyonlands National Park, in¬ 
cluding Myotis califomicus, M. evotis, M. thysanodes, 
M. yumanensis, Lasiurus cinereus, Parastrellus hes- 
perus, Eptesicus fuscus, Corynorhinus townsendii, 
Idionycteris phyllotis, and Antrozous pallidus. 

Schafer (1991) reported the occurrence of six spe¬ 
cies of bats from the Abajo Mountains ( Myotis evotis, 
M. ciliolabrum, M. volans, Lasionycteris noctivagans, 
Eptesicus fuscus, and Lasiurus cinereus ). Mollhagen 
and Bogan (1997) documented 15 species of bats 
from the Henry Mountains and surrounding areas in 
southeastern Utah, including those reported by Schafer 
(1991) plus Euderma maculatum and Tadarida brasil- 
iensis; and Mollhagen and Bogan noted the nearby 
capture of Nyctinomops macrotis. Bogan et al. (2006), 
working in Canyonlands National Park, reported 16 
species, excluding Myotis lucifugus, records of which 
are problematic in the region. 

Although long-lived (5-20 yrs), bats as a group 
are of concern because they have low reproductive 
output (usually 1 young/female/yr) and may roost in 
large groups (hundreds to thousands) where they are 
susceptible to human disturbance (O’Shea and Bogan 
2004). Furthermore, much of the existing data on chi¬ 
ropteran population trends is of limited use; because of 
methodological issues with collecting the data, limited 
time periods of data collection, or inadequate sample 
sizes for statistical analysis (O’Shea and Bogan 2004). 

Our objectives were to: (1) conduct an inventory 
of all bat species in Arch Canyon; and (2) to identify 
differences in occurrences of bats, if any, between upper 
and lower reaches of the canyon. 


Methods 


The study area of the bat survey was the portion of 
Arch Canyon between the 4,977 ft (1,517 m) and 5,621 
(1,713) elevations. The coordinates are N37.545079 
x W109.666013 and 37.60657 x 109.761735, respec¬ 
tively. The lower boundary is marked by a cattle- 
guard, a fence and a visitor registration station. The 
upper boundary lacks a fence but, at the time of our 


work, there were Forest Service markers delineating 
the boundary line. One of the two state sections of a 
16-section township extends across the study area. The 
state section divides the canyon into nearly equal upper 
and lower portions. There are no obvious markers, but 
this reach of the canyon lies approximately between 
the coordinates of N37.563017 x W109.719205 and 


Mollhagen and Bogan—Inventory of Bats in Arch Canyon, Utah 


217 


37.566902 x 109.730099. No work was undertaken 
on the state section. 

We were defeated by extreme flooding on our ini¬ 
tial visit to Arch Canyon in September 2006. We could 
not pass the second crossing and mist nets yielded no 
captures. In June, 2007, there was again considerable 
water, but we were able to identify 19 sites with pools 
over which we might capture bats. We mist netted on 
six consecutive nights in the period 12-17 June. We 
netted again on six consecutive nights 11-16 July. 
Another flash flood had occurred in the canyon a few 
days prior to our arrival on 6 August 2007. We hiked to 
our previously-utilized netting sites on 7 and 8 August. 
After some trail repair on 9 August, we were able to 
drive approximately 1.5 mi (2.4 km) up the canyon. 
Stream flow was nearly equivalent with the previous 
September. We did not continue the netting effort in 


the upper canyon because of poor netting success, the 
abundance of muddy water in the canyon, the onset of 
the monsoon rain season, and the poor state of the trail. 

We recorded GPS coordinates and elevations for 
each netting site using Garmin GPS units set to record 
coordinates in decimal degrees (WGS84 datum; Table 
1). Elevations were reconciled with USGS quad maps 
and, when there was a discrepancy between sources, 
interpolated values were used. The distance from the 
entrance to each netting location was also documented. 
We made daily journal entries while in the field, and 
recorded all data onto pre-printed datasheets. The 
data were later entered into an Excel spreadsheet for 
analysis. 

We believe that quiet, permanent pools with 
fetches of at least 25 ft (8 m) best serve the continuum 


Table 1. Locations in Arch Canyon, Utah. Locations listed are the USFS boundary at the upper part of the 
study area (USFS), the five sampling locations in the upper canyon (U5 to Ul), the upper boundary of the state 
section (State), the lower boundary of the state section (Section), the five sampling locations in the lower canyon 
(L5 to LI), and the entrance to the canyon (Entrance). In the right-most column are the trail miles to each of 
the landmark locations. Data presented are coordinates as decimal degrees (WGS84 datum) and elevation in 
both feet and meters. 


Northing 

Westing 

Elevation (ft) 

Elevation (m) 

Trail Miles 

USFS 

37.606570 

109.761735 


8.9 

U5 

37.605416 

109.760757 

5598 

1707 

00 

bo 

U4 

37.604525 

109.759209 

5596 

1707 

8.7 

U3 

37.578576 

109.737058 

5396 

1646 

6.1 

U2 

37.576471 

109.734863 

5374 

1639 

5.7 

Ul 

37.568405 

109.730730 

5312 

1620 

5.1 

State 

37.566902 

109.730099 


5.0 

Section 

37.563017 

109.719205 


4.2 

L5 

37.561455 

109.718088 

5270 

1607 

4.1 

L4 

37.550217 

109.683029 

5045 

1539 

1.5 

L3 

37.549276 

109.679446 

5038 

1537 

1.3 

L2 

37.547641 

109.676701 

5024 

1532 

1.1 

LI 

37.546128 

109.670412 

4833 

1474 

0.7 

Entrance 

37.545079 

109.666013 


0.0 


218 


Clyde Jones Memorial Volume 


between the small, slow, agile bat species and the 
large, swift, direct-flying species. We were seeking 
locations that exhibited some evidence of permanence 
so we might return to these sites later in the season. 
This evidence included an apparent spring discharge, 
water depth of a foot or more, fishes larger than fry 
in the pools, and the occurrence of obligate wetlands 
vegetation. Many of these sites occur at trail crossings, 
but it was not clear if the pools were created by vehicle 
traffic, or if the pool margins simply provided gentle 
slopes for the passage of vehicles. 

Our main objective was to compare faunas above 
and below the state section of Arch Canyon. Pools 
L1-L5 (Table 1) are in the reach below the state sec¬ 
tion, whereas U1-U5 are above the state section. LI 
is the lower-most pool whereas U5 is the upper-most 
site. We found many more possible netting locations 
on our first reconnaissance, but the aforementioned 
localities were the only 10 that met our requirements 
for the duration of the project. 

We initially prepared a list of bats for Arch Can¬ 
yon that included all species that might occur in the 
region. Primary references for this list were Durrant 
(1952), Hall (1981), Armstrong (1982), Mollhagen and 
Bogan (1997), Haymond et al. (2003), Bogan et al. 
(2006) and a series of unpublished reports by Bogan 
and cooperators who have inventoried mammals oc¬ 
curring in southern Utah national parks. This original 
list included 16 species, not including M. lucifugus. 

Mist nets were deployed across and around bodies 
of water to capture bats flying in to drink or foraging on 
insects over the water. Mist netting especially is effec¬ 


tive when sources of water in the landscape are limited, 
as this causes bats to be concentrated in a relatively 
small area where they are more susceptible to capture. 

The lengths of nets ranged from 9 to 60 ft (3-20 
m) and the numbers of nets deployed on any single eve¬ 
ning varied from one to five, depending upon the pool 
surface area and shape. Mist nets were set up shortly 
before sunset and tended for several hours until activity 
declined; in some cases nets were observed throughout 
the night. The nets were never left untended. 

Bats were immediately removed from nets fol¬ 
lowing capture. The time of capture, species, sex, 
reproductive condition, and any miscellaneous com¬ 
ments were recorded on standardized data sheets. 
The unharmed bats were released within minutes after 
capture in the net. All participants in this work were 
experienced and knowledgeable in capturing, han¬ 
dling, and identifying bats. Personnel handling bats 
previously had taken the standard pre-exposure rabies 
immunization and had demonstrated titers indicating 
rabies antibody activity. 

Common and scientific names of mammals fol¬ 
low those of Bradley et al. (2014). All capture and 
handling of animals was performed in accordance 
with the written protocols approved by the American 
Society of Mammalogists. Some individuals of previ¬ 
ously undocumented species from the study area were 
retained and prepared as voucher specimens (skins 
and skeletons) and deposited in the USGS Biological 
Surveys Collection in the Museum of Southwestern 
Biology at the University of New Mexico. No speci¬ 
mens of species of concern were prepared. 


Results 


We netted at five sites both above and below 
the state section (Table 1). We captured a total of 295 
individual bats of 15 species at the 10 collecting sites 
(Table 2). The total number of 15 species obtained in 
this study is consistent with the results of surveys in 
the Henry Mountains by Mollhagen and Bogan (1997) 
and in Canyonlands National Park (Bogan et al. 2006). 


The eight most common species of bats were 
Parastrellus hesperus (32.9% of total captures), Nyc- 
tinomops macrotis (23.4%), Tadarida brasiliensis 
(12.5%), Eptesicus fuscus (7.1%), Antrozous pallidus 
(5.4%), Myotis yumanensis (4.7%), Euderma macula- 
tum (3.7%), and M. californicus (3.4%). We captured 
1-6 individuals (less than 3% of total captures) for 


Mollhagen and Bogan—Inventory of Bats in Arch Canyon, Utah 


219 


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220 


Clyde Jones Memorial Volume 


each the seven other species. By month, we captured 
93 individuals and 10 species in June, 191 individuals 
and 10 species in July, and 11 individuals and 3 species 
in a storm-shortened August trip. We saw very few bats 
flying in the canyon in August. 

Above the state section we captured a total of 
159 individuals and 15 species (Table 2); below the 
state section, we took 136 of 11 species. Those species 
captured only above the state section were M. evotis, 
M. thysanodes, M volans, and E. maculatum. Most 
species were caught in roughly equal numbers above 
and below the state section, although slightly more P. 
hesperus (52 above, 34 below) and significantly more 
A. pallidus were captured above the state section (13:1). 
More Free-tailed Bats ( T. brasiliensis and N. macrotis) 
were captured in the lower reach of the canyon (58) 
than in the upper (43). 


Given the relatively short study period we are 
reluctant to emphasize these differences too much. 
However, our impressions of habitat preferences 
of these bats gained from this (and previous) study 
are that those four species caught only in the upper 
canyon may prefer somewhat higher elevations; espe¬ 
cially M. evotis and M. volans. Euderma maculatum 
is known from a variety of habitats but the species 
may be somewhat more common at upper elevations. 
However, the slightly increased capture rates of two 
arid-adapted species, A. pallidus and P. hesperus, at 
the upper sites suggests that upper Arch Canyon may 
possess some subtle advantage for some species. We 
are reluctant to emphasize this because there seemed 
to be a greater number of larger pools, in the lower 
canyon, particularly in the lower reaches. The greater 
availability of water at the lower elevation netting sites 
may have rendered netting efforts less effective than 
in the upper canyon. 


Discussion 


We suspect that most of the species we captured 
have roosts in the canyon and many probably hiber¬ 
nate nearby. The two species of tree-roosting bats (L. 
cinereus and L. noctivagans ) occur seasonally and 
presumably migrate to the south prior to winter. Most 
individuals of these two species in the area are males. 
Females occur farther east, where they give birth 
(Cryan 2003). Tadarida brasiliensis and N. macrotis 
are likewise believed to be migratory, spending the 
winter to the south. We are aware of one small and one 
large maternity colony for N. macrotis along nearby 
Comb Wash. 

Armstrong (1982) postulated two additional spe¬ 
cies to occur in the general area of Arch Canyon, the 
Little Brown Myotis (M lucifugus ) and the Eastern Red 
Bat ( Lasiurus borealis ). The former is represented by 
a specimen captured at French Spring in Glen Canyon 
National Recreation Area (GCNRA). We have exam¬ 
ined the specimen (University of Colorado Museum 
no. 15207) and it is instead a M. volans. However, M. 
lucifugus does occur in southern Utah. We have exam¬ 
ined specimens from the Fremont River in the vicinity 
of Bicknell and Donkey Lake on Boulder Mountain, 
Wayne County, Utah (specimens in Utah Museum of 
Natural History). Hasenyager (1980) listed two oc¬ 


currences of M. lucifugus from Bluff City, San Juan 
County but we have found no specimens to document 
these records during our travels to Utah museums. 
As Oliver (2000) noted, recent studies of bats across 
southern Utah have failed to document the occurrence 
of M. lucifugus, and it seems parsimonious to assume 
the species is absent from much of this area. Should 
a short-eared, large-footed, medium-sized myotis be 
captured that is not M. yumanensis, it should be exam¬ 
ined carefully as it may actually be Myotis occultus. 

Armstrong (1982) also listed the possible oc¬ 
currence of Lasiurus borealis in the area. He referred 
to the records of Hardy (1941) from Carbon County 
(presumably Kenilworth Mine, Price) and from Wash¬ 
ington County (La Verkin Cave; St. George). We 
searched for the origins of these records and located 
only a specimen from the Kenilworth Mine at the 
museum at California State University, Long Beach. 
Our supposition is that the specimens were in Hardy’s 
personal collection and were taken to Long Beach when 
Hardy left employment at what was then Dixie Junior 
College. We have not examined this specimen as of 
this writing and are uncertain whether the Kenilworth 
specimen represents the Eastern Red Bat (L. borealis ) 
or Western Red Bat (L. blossevillii ) as now recognized 


Mollhagen and Bogan—Inventory of Bats in Arch Canyon, Utah 


221 


(Oliver 2000). Although the Carbon County record 
could have been L. borealis , the absence of any records 
of this species since 1937 from anywhere in eastern 
Utah suggests this species should not be considered as 
part of the regional bat fauna. The nearest records are 
in northeastern Colorado. Conversely, there continue 
to be reports of L. blossevillii from southwestern Utah 
and a young-of-the-year male from Springville, Utah 
County (Mollhagen and Bogan 1997; Oliver 2000). 
Nonetheless, we have engaged in considerable field 
work where L. blossevillii might seasonally inhabit 
(Kane, Garfield, Wayne and San Juan counties) and 
none have been captured. Thus, we regard this species 
as of unlikely occurrence in the Arch Canyon area. 

We did not catch Myotis ciliolabrum in Arch 
Canyon, although we are confident the species occurs 
in the area, at least seasonally. This species likely is 
restricted to higher elevation areas of southeastern Utah 
for most of the year. They have been captured in the 
Henry Mountains (Mollhagen and Bogan 1997). The 
species was not captured in the Canyonlands National 
Park (CANY) survey (Bogan et al. 2006), but it was 
recorded acoustically. 


The Utah bat species of concern are M. thy- 
sanodes, I. phyllotis, C. townsendii, E. maculatum and, 
N. macrotis. A comparison of the occurrence of these 
species in Arch Canyon with other bat inventories on 
the Colorado Plateau is warranted. There are six such 
inventories. The Henry Mountains and Canyonlands 
National Park inventories presented in Table 3 are 
within 100 miles of Arch Canyon. The other four inven¬ 
tory sites are greater distances away in New Mexico, 
Arizona, Colorado and Utah. 

Mollhagen and Bogan (1997) captured bats from 
22 separate localities in the Henry Mountains (Table 
3). The Henrys are approximately 80 miles west and 
differ from Arch Canyon in occupying a much larger 
geographic area, having a greater elevational relief and, 
consequently, much greater habitat variation. Bogan 
et al. (2006) conducted a two-year study of bats in 
Canyonlands National Park (CANY), approximately 
50 miles to the north. Bats were captured at 32 locali¬ 
ties over a wider area and a greater elevational range 
than occur in Arch Canyon. Both of these studies had 
many more total captures (Table 3), nearly double in 
the Henry Mountains and over five times the number 
in CANY. 


Table 3. Comparison of the abundance of Utah bat species of concern among six inventories for bats on the Colorado 
Plateau. The studies are as follows: Arch Canyon, UT, this report; Henry Mountains, UT, Mollhagen and Bogan 
(1997); Jemez Mountains, NM, Bogan et al. (1998); Coconino Plateau, AZ, Rabe et el. (1998); Morell et al. (1999); 
Canyonlands National Park, UT, Bogan et al. (2006); Dinosaur National Monument, Bogan and Mollhagen (2010); 
and Mesa Verde National Park, O’Shea et al. (2011). Data shown are the total number of captures for each study and 
the respective percentage of the total fauna of each species of concern. The acronyms for the bat species are as fol¬ 
lows: Myth, Myotis thysanodes ; Idph, Idionycteris phyllotis ; Coto, Corynorhinus townsendii ; Euma, Euderma macu¬ 
latum ; and Nyma, Nyctinomops macrotis. 


Species 


State(s), region, and total bat captures for each study 


UT, 

Henry 

Mountains, 

572 

UT, 

Canyonlands 

NP, 

1,717 

CO, 

Mesa Verde 
NP, 

1,996 

UT, CO, 
Dinosaur NM, 
909 

AZ, 

Coconino 

Plateau, 

1,532 

NM, 

Jemez 

Mountains, 

1,532 

UT, 

Arch 

Canyon, 

295 

Myth 

34 (5.9%) 

64 (3.7%) 

41 (0.1%) 

7(1.5%) 

122 (7.2%) 

69 (4.5%) 

1 (0.3%) 

Idph 

9(1.6%) 

36(2.1%) 

0 

0 

26(1.5%) 

0 

4(1.4%) 

Coto 

16(2.8%) 

23 (1.3%) 

13(<0.1%) 

14 (3.0%) 

2 (< 0.1%) 

7 (0.5%) 

2 (0.7%) 

Euma 

1 (0.2%) 

0 

10(0.1%) 

5(1.1%) 

0 

12 (0.8%) 

11 (3.7%) 

Nyma 

0 

3 (0.2%) 

0 

0 

0 

15(1.0%) 

69 (23.3%) 


222 


Clyde Jones Memorial Volume 


Even though the other four other bat inventories 
(Table 3) are hundreds of miles away, their respective 
faunas each have at least three of the five Utah bat spe¬ 
cies of concern. In northern New Mexico, Bogan et 
al. (1998) captured 1,532 bats in the Jemez Mountains, 
at Bandelier National Monument and the Los Alamos 
National Laboratory. There are two reports on the same 
bat fauna (1,684 captures) on the Coconino Plateau in 
northern Arizona, Rabe et al. (1998) and Morrell et al. 
(1999). Bogan and Mollhagen (2010) reported on a 
reinventory of the bats in Dinosaur National Monu¬ 
ment. They captured 909 bats at localities in north¬ 
western Colorado and northeastern Utah. O’Shea et al. 
(2011) detailed an inventory of 1,996 bats captured in 
southwestern Colorado, in Mesa Verde National Park. 

In a direct comparison among the faunas, M. 
thysanodes seems underrepresented in Arch Canyon 
and Mesa Verde NP (Table 3); we captured only one 
individual in Arch Canyon. Only four individuals of I. 
phyllotis were captured in Arch Canyon, but there is no 
material difference in the percent representation among 
the four faunas where Idionycteris is represented. 
Based on the brevity of this study and small sample 
size, C. townsendii may also be underrepresented in 
Arch Canyon. 

However, 11 captures of E. maculatum seem 
remarkable (Table 3). To put this into perspective, 
fieldwork in the Henry Mountains continued well 
beyond the published study period, until 2003. This 


We appreciate the assistance of Tammy Wallace 
and Scott Edwards of BLM in sharing their knowledge 
of the canyons and the water sources. Cindy Ramotnik, 
USGS, assisted in processing the voucher specimens in 
the museum. Trish Freeman and Cliff Lemen assisted 
in the field during the abortive August 2007 trip. Frank 


extension of sampling time more than doubled the total 
bat captures (unpublished data), but no additional Eud¬ 
erma were captured. This would reduce the percent of 
Spotted Bat captures to approximately 0.1. Although no 
Eudenna were captured in Canyonlands, the vocaliza¬ 
tions of two individuals were heard over the course of 
that study. Including these records in the calculation 
of the percent of fauna at Canyonlands also yields a 
value of approximately 0.1. The percents of Euderma 
captures are similar among the other three studies. 

The capture of Nyctinomops is not surprising to 
us because there are two known maternity colonies in 
nearby Comb Ridge, and we had captured Nyctinomops 
in Arch Canyon some years ago (also see Zimmerman 
1970). However, the capture of 69 individuals, rep¬ 
resenting over 23 percent of the total sample of Arch 
Canyon, is unusually high in comparison with the 
Henry Mountains, Canyonlands, and Jemez Mountains 
bat faunas (Table 3). No N. macrotis had been taken 
in the Henry Mountains at the time of the Mollhagen 
and Bogan (1997) report. Subsequent field work, more 
than doubling the total bats captured, has produced only 
a single individual of this species (unpublished data). 
Thus Nyctinomops ’ contribution to the total fauna is 
approximately 0.1 percent. The respective value in 
Canyonlands is 0.2 percent. Clearly, Euderma and 
Nyctinomops were common at the time of our study, 
perhaps because Arch Canyon and vicinity provided 
abundant roosting sites, foraging sites, and water. 


Montella of Blanding also rendered towing assistance 
during the same August trip. Finally we are grateful for 
the skillful reviews by Jim Goetze and Rick Manning, 
whose efforts contributed mightily to the final form of 
the manuscript. 


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Addresses of authors: 

Tony R. Mollhagen 

4524 64th Street 
Lubbock, TX 
shellbadger@yahoo.com 


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Michael A. Bogan 

PO Box 2452 
Corrales, NM 87048 
mbogan @ unm. edu 


Comparison of Pasturelands Containing Texas Kangaroo Rat (Dipodomys 
elator) Burrows to Adjacent Roadsides in Wichita County, Texas, with 
Comments on Road Usage by D. elator 

Jim R. Goetze, Allan D. Nelson , and Larry L. Choate 


Abstract 

The Texas Kangaroo Rat ( Dipodomys elator) is a threatened species in Texas. This 
heteromyid has been nominated for federal protection under the Endangered Species 
Act. The United States Fish and Wildlife Service completed a 90-day finding on the 
petition and stated that the use and effects of roads on the Texas Kangaroo Rat was un¬ 
clear. We surveyed pasturelands and adjacent roadsides for D. elator in Wichita County, 
Texas, and examined differences in vegetation composition between pasturelands and 
roadsides. Over 1,000 trapnights were conducted in each habitat. Only three percent 
of D. elator captures occurred along roadsides. Roadside vegetation was taller, had less 
bare ground, more introduced grass species, and higher species richness than adjacent 
pasturelands. Forb coverage was similar between the pasturelands and roadsides. In 
Wichita County, D. elator uses pastureland more frequently than adjacent roadsides. 
This is likely due to differences in vegetation between the two habitats. 

Key words: Dipodomys elator , habitats, pastureland, roadside, Texas, Texas 
Kangaroo Rat, vegetation, Wichita County 


Introduction 


The Texas Kangaroo Rat, Dipodomys elator, 
is unusual because the habitat in which it is found is 
not typical for kangaroo rats. It seems to prefer soils 
with high clay content which support overgrazed or 
short grasses (Dalquest and Collier 1964; Roberts and 
Packard 1973; Dalquest and Homer 1984; Stangl et al. 
1992; Schmidly 2004; Goetze et al. 2007; Nelson et al. 
2009; Stasey et al. 2010) and has rarely been recorded 
in locations with dense vegetation. 

As a result of overgrazing and control of wildfires, 
mesquite ( Prosopis glandulosa ) and other disturbance- 
related shrubs, grasses and forbs have increased in 
abundance across much of Wichita County, Texas, and 
habitat modification, such as conversion of pastureland 
to monoculture, has resulted in extensive fragmentation 
of Texas Kangaroo Rat habitat (Diamond and Shaw 
1990). From 1996 to 2000, Martin (2002) surveyed the 


historic range of D. elator and found the range of D. 
elator reduced from 10 to five counties in north Texas. 
He did not find D. elator in Oklahoma (Martin 2002). 
This concurs with other researchers who have been un¬ 
able to locate any populations ofD. elator in Oklahoma 
(Jones et al. 1988; Moss and Mehlhop-Cifelli 1990). 

Federally, D. elator was listed as a Category 2 
candidate species under the Endangered Species Act 
of 1973 (Martin 2002). Category 2 candidates were 
formerly considered species of concern for the United 
States Fish and Wildlife Service (USFWS), and en¬ 
dangered or threatened status was possibly warranted. 
However, insufficient data existed to justify an elevated 
listing (USFWS 1996). 

In Texas, D. elator is listed as a threatened species 
by the Texas Parks and Wildlife Department (TPWD) 


225 


226 


Clyde Jones Memorial Volume 


(Martin 2002; Schmidly 2004). Reasons for D. ela¬ 
tor being listed as a threatened species by the TPWD 
are based on scarcity of this species and the small 
geographic range from which it is known (Stangl and 
Schafer 1990). 

Most recently, WildEarth Guardians petitioned 
USFWS to federally list the Texas Kangaroo Rat 
(WildEarth Guardians 2010). In the petition to 
USFWS, WildEarth Guardians stated that many factors 
had contributed to the decline of D. elator across its 
historic range. Among the factors WildEarth Guardians 
listed were: 1) loss of native habitat to cropland; 
2) loss of historic ecological processes involving 
bison, prairie dogs and fire; 3) domestic livestock 
grazing; 4) development and roads; 5) brush control; 
6) overharvesting for scientific purposes; 7) parasites 
and predation; 8) little regulatory protection; and 9) 
climate change. The USFWS responded with a 90-day 
finding that concluded the petition presents substantial 
scientific information indicating that listing the Texas 
Kangaroo Rat throughout its entire range may be 
warranted (USFWS 2011). This finding was based on 
the present or threatened destruction, modification, or 
reduction of the Texas Kangaroo Rat’s habitat or range, 
and the inadequacy of existing regulatory mechanisms 
(USFWS 2011). Specifically, the USFWS found that 
the loss of burrowing habitat and genetic isolation of 
populations due to the conversion of native rangeland 
to agricultural cropland, and the inadequacy of existing 
regulatory mechanisms to protect against such land 
conversion, may pose a threat to the Texas Kangaroo 
Rat throughout all or a significant portion of its range 
(USFWS 2011). We agree with the assessment of the 
USFWS, and within this paper expand upon vegetation 
associated with roads and adjacent pasturelands in 
Wichita County and presence of D. elator. 


Much of Martin’s research methodology involved 
roadside surveys and trapping along roadsides (Martin 
2002). Other researchers have studied the effects of 
roads as potential corridors or barriers to distribution 
and movement of small mammals, including species of 
kangaroo rats (Roberts and Packard 1973; Brock and 
Kelt 2004). Stephen’s Kangaroo Rat ( D . stephensi ) 
is known to colonize dirt roadsides (O’Farrell 1990), 
and Brock and Kelt (2004) noted that D. stephensi may 
occur in small, linear populations along roadsides even 
when adjacent habitats are unsuitable. 

Although a relatively large number of studies 
have examined aspects of their ecology (Dalquest 
and Collier 1964; Roberts and Packard 1973; Jones et 
al. 1988; Moss and Mehlhop-Cifelli 1990; Stangl and 
Schafer 1990; Stangl et al. 1992; Martin 2002; Stasey 
2005; Goetze et al. 2007; Goetze et al. 2008; Nelson 
et al. 2009; Stasey et al. 2010) and systematics (Mer- 
riam 1894; Johnson and Selander 1971; Hamilton et al. 
1987; Dalquest et al. 1992; Mantooth et al. 2000), few 
investigators have closely examined vegetation associ¬ 
ated with D. elator burrows (Martin and Matocha 1991; 
Martin 2002; Nelson et al. 2009). No researchers have 
specifically examined roadside vegetation adjacent 
to pasturelands containing known populations of the 
Texas Kangaroo Rat. 

Because previous researchers have utilized road 
surveys to verify the presence of D. elator within its 
current range, the primary goals of this investigation 
were to quantify vegetative characteristics within 
pasturelands that contained burrows where the Texas 
Kangaroo Rat was obtained and to gather preliminary 
information concerning suitability of roadsides as 
habitat for D. elator within Wichita County, Texas. 


Methods 


Twenty-one active D. elator burrows were identi¬ 
fied from Wichita County from four different localities 
(Fig. 1). Trapping to test for burrow occupancy was 
conducted by placing three 7.5 x 8.8 x 30 cm Sher¬ 
man Five Traps within 0.10 to 0.50 m of each burrow 
entrance, with the open end of each trap facing the en¬ 
trance (Cross and Waser 2000). Traps were baited with 
dry oatmeal each evening and checked each morning. 


If D. elator were captured, vegetation was sam¬ 
pled during May using aim 2 quadrat placed directly 
over burrows where D. elator were captured. Burrow 
parameters were taken in May to avoid seasonal influ¬ 
ences. Within each quadrat, vegetative richness was 
recorded as total number of species present. Percent¬ 
age coverage of grass, forbs, and bare ground within 
each quadrat was recorded, as was average herbaceous 


Goetze et al.—Pasture and Roadside Use by Dipodomys elator 


111 


N 


FM 1739 


m 

Site D 


/ 


Site A 

SiteC 


Goetze Road 


T 


& 

Site B 

Burnett Ranch Road 


Hall Road 


4 km 


Lake Buffalo Creek Reservoir 


Figure 1. Map showing the four sampling locations in Wichita County with associated roads 
where study samples and data were taken. Geographic coordinates to the entrance gates of 
the four localities are Site A (34.05756N, 98.69716W), Site B (34.06497N, 98.70709W), Site 
C (35.05446N, 98.78699W), and Site D (34.05423N, 98.781721W). 


vegetation height (obtained by averaging the height of 
the herbaceous vegetation 15 cm interior to each comer 
of the quadrat). Specimens of the dominant herbaceous 
plants were collected, identified with appropriate keys 
(Diggs et al. 1999), and made into vegetation vouchers 
subsequently deposited in the herbarium of Tarleton 
State University. 

During the course of our study, no Texas Kan¬ 
garoo Rat burrows were tentatively identified along 
roadsides. Therefore, a 100 m tape was placed along 
roadside traplines and random numbers were selected 
as sites for quadrat sampling as described above. A 


total of 20 quadrats were sampled along roadsides 
adjacent to the aforementioned pastureland localities 
(Fig. 1). To survey for the presence of the Texas Kan¬ 
garoo Rat in the absence of burrow evidence, Sherman 
live traps were placed approximately 10 m apart in a 
transect along roadside fences, baited, and checked as 
described above. 

Averages and ranges for vegetative data as well 
as comparisons between sites were calculated in Mi¬ 
crosoft Excel. Comparisons between pasturelands 
and adjacent roadsides were made using SigmaPlot 12 
software (Systat 2016) to calculate a Mann-Whitney 


228 


Clyde Jones Memorial Volume 


Rank Sum Test. Manifold 8.0 GIS software program 
(Manifold Software Limited 2013) and a digital, 1 m 
resolution, orthographic quadrangle (DOQ) map of 


Wichita County from the Natural Resources Conserva¬ 
tion Service (NRCS) were utilized to generate the base 
map for Figure one. 


Results 


In May of 2011, 2012, and 2015 we character¬ 
ized the vegetation associated with 21 Texas kangaroo 
Rat burrows from four locations in Wichita County as 
well as 20 quadrats along roadsides adjacent to the four 
localities. During this period of time, trapping efforts 
consisted of a total of 1,031 trap nights within pasture- 
lands and 1,045 trap nights along adjacent roadsides. 

The most common plant dominants in the pas- 
turelands were Little Barley (Hordeum puscillum ) and 
Virginia Pepper-grass (Lepidium virginica) (Table 1). 
Next in vegetation dominance were Whorled Dropseed 
(Sporobolus pyramidatus ) and Silverleaf Nightshade 
(Solatium eleagnifolium). The most common domi¬ 
nants along the roadsides were Johnsongrass (Sorghum 
halepense ) and an unknown forb (Table 1). During 
sampling time, the unknown forb consisted of only a 
short stem with a few leaves and was unidentifiable. 
Next in vegetation dominance were two introduced 
bromes (Bromus catharticus and B.japonica ) and two 
native species, Needlegrass (Nasella leuchotricha) and 
Western Ragweed (Ambrosia psilostachys). Average 
herbaceous height, percent bare ground, percent grass¬ 


es, and species richness were significantly different 
between pasturelands and adjacent roadsides (Table 1). 
Average percent forbs was not significantly different (p 
= 0.07). Averages for all of the vegetation parameters 
were greater in the roadsides except for percent bare 
ground, which was greater in the pasturelands (Table 1). 

Along with D. elator, five species of mammals 
were captured in the pastureland sites, whereas three 
species were obtained along the adjacent roadsides 
(Table 2). The Texas Kangaroo Rat was the most com¬ 
monly obtained species within pasturelands and the 
White-footed Deermouse (Peromyscus leucopus ) was 
the most commonly captured mammal along roadsides. 
Within the pastureland sites, both P. leucopus and the 
North American Deermouse (P. maniculatus ) were 
obtained, but no P. maniculatus was captured along 
roadsides during the study. The Hispid Cotton Rat 
(Sigmodon hispidus ) and Hispid Pocket Mouse (Chae- 
todipus hispidus ) were obtained in both pasturelands 
and roadsides, whereas the Southern Plains Woodrat 
(Neotoma leucodon ) was captured only in pasturelands 
(Table 2). 


Table 1. Dominant vegetation and average herbaceous height, percent coverage of bare ground, 
forbs, grasses, and species richness in the pastureland and adjacent roadsides. Ranges are en¬ 
closed in parentheses following means. An asterisk indicates significant differences between 
means at the < 0.001-0.01 levels. 


Pastureland 
(* = 21) 

Roadside 
(n = 20) 

Dominant forb 

Virginia Pepperweed 

Unknown 

Dominant grass 

Little Barley 

Johnsongrass 

♦Average herbaceous height (cm) 

13.2 (0.0-59.0) 

69.7 (6.9-129.3) 

♦Average % bare ground 

51.7(20.0-94.0) 

11.2(0.0-20.0) 

Average % forbs 

11.9(0.0-50.0) 

19.8(1.0-65.0) 

♦Average % grasses 

25.0(1.0-45.0) 

69.0 (10.0-98.0) 

♦Average richness 

5.0(1.0-12.0) 

6.6 (4.0-10.0) 


Goetze et al.—Pasture and Roadside Use by Dipodomys elator 


229 


Table 2. Mammalian species and numbers obtained from live traps in pasturelands and adjacent 
roadsides. 


Species 

Pastureland 

Roadside 

Dipodomys elator 

36 

1 

Peromyscus maniculatus 

17 

0 

Peromyscus leucopus 

14 

29 

Sigmodon hispidus 

2 

3 

Neotoma leucodon 

2 

0 

Chaetodipus hispidus 

4 

2 


Discussion 


WildEarth Guardians (2010) suggested that all 
roads within the range of D. elator increased the risks 
of predation and mortality from vehicle collisions. 
USFWS (2011) indicated that Texas Kangaroo Rats 
(Roberts and Packard 1973; Stangl and Schafer 1990; 
Stangl et al. 1992), and other species of kangaroo rats 
(Brock and Kelt 2004), may preferentially use dirt roads 
as migration corridors. Although there are reports of 
specimens killed by vehicular traffic (Dalquest and 
Collier 1964; Jones et al. 1988), USFWS suggested 
that roads were not having a negative impact on Texas 
Kangaroo Rat mortality. During the course of our study, 
only a single D. elator was salvaged from a paved 
roadside adjacent to Focality A (Fig. 1). 

USFWS (2011) also mentioned that it is well 
established that nighttime road surveys are an easy and 
effective way to determine the presence of the Texas 
Kangaroo Rat, suggesting the Texas Kangaroo Rat 
does not entirely avoid these areas. During our study, 
we captured only one of 37 D. elator from a roadside 
location (site D; Fig. 1). When released at the roadside, 
this individual entered a burrow within the adjacent 
pastureland. All D. elator burrows were found within 
pasturelands. Our results differ from those of Stapp 
and Findquist (2007) who worked with D. ordii. In 
a study of Ord’s Kangaroo Rat in Colorado, Stapp 
and Findquist (2007) reported greater numbers of D. 
ordii captured from roadsides as opposed to adjacent 
pasturelands. However, the vegetation in the Colorado 
pasturelands was similar to the cover and height of our 
Wichita County roadside vegetation. Brock and Kelt 


(2004) reported usage patterns similar to D. elator for 
Stephen’s Kangaroo Rat in California. 

Based on the results of our study, vegetation and 
numbers of Texas Kangaroo Rats found in pasturelands 
and along adjacent roadsides often significantly differ 
(Tables 1 and 2). Brock and Kelt (2004) and Stapp and 
Findquist (2007) reported significant use of roadsides 
by D. stephensi and D. ordii for burrowing and foraging 
sites. In our study, 37 D. elator were captured within 
pasturelands and only one Texas Kangaroo Rat was 
captured along a roadside. However, four of the 21 
burrows within pasturelands were along unimproved 
(dirt) roads. This supports the findings of Brock and 
Kelt (2004) regarding Stephen’s Kangaroo Rat and 
Stapp and Findquist (2007) regarding Ord’s Kangaroo 
Rat. The Wichita County roads sampled adjacent to the 
pasturelands were improved roads (gravel or asphalt). 
Although Roberts and Packard (1973) stated that D. 
elator used roads as burrow locations, we did not find 
that to be the case in our study. 

Dominant vegetation found at burrows within the 
pasturelands was mostly native. Fittle Barley was the 
most dominant native grass with Whorled Dropseed 
being the second most common native grass. Intro¬ 
duced grasses dominated in roadsides and included 
Johnsongrass, Japanese Brome, and Rescue Grass. 
As we surveyed for trapping locations, we often en¬ 
countered dense concentrations of introduced grasses 
such as Japanese Brome. Similarly, introduced plant 
numbers were higher along roadsides than in grasslands 


230 


Clyde Jones Memorial Volume 


in the D. ordii study in Colorado (Stapp and Lindquist 
2007). Increases of these introduced grasses likely 
will negatively affect the Texas Kangaroo Rat and are 
probably why many sites, especially along roadsides, 
no longer have Texas Kangaroo Rats. Dense vegeta¬ 
tion likely impedes burrow construction and prevents 


Texas Kangaroo Rats from locating potential predators. 
In addition, dense vegetation reduces bare patches 
needed for dust bathing (Goetze et al. 2008). In Wichita 
County, D. elator uses pastureland more frequently than 
adjacent roadsides. This is likely due to differences in 
vegetation between the two habitats. 


Acknowledgments 


We thank the Goetze family for use of their 
property. We also thank Texas Parks and Wildlife De¬ 
partment (TPWD) and United States Fish and Wildlife 
Service for partial funding under TPWD Contract No. 


403892. We also wish to thank T. R. Simpson and 
other reviewers for offering comments to improve the 
manuscript. 


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Addresses of authors: 

Jim R. Goetze 

Department of Natural Sciences and Kinesiology 
Laredo Community College 
Laredo, TX 78040 
j goetze @ laredo. edu 


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{Dipodomys elator ): Biological habits and population 
estimation. Unpublished Masters Thesis. Tarleton State 
University, 45 pp. 

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2010. Differential use of grazed and ungrazed plots by 
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Central Texas. Texas Journal of Science 62:3-14. 

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cies Act. WildEarth Guardians, Denver, Colorado. 32 pp. 


Larry L. Choate 

Department of Biology (Emeritus) 
East Central University 
Ada, OK 74820 
Ichoate @ ecok. edu 


Allan D. Nelson 

Department of Biological Sciences 
Tarleton State University 
Stephenville, TX 76402 
nelson @ tarleton. edu 


Encomia and Reflections 


Don Wilson (p. 49) 


Clyde Jones was a people person. He had a knack 
for making everyone feel comfortable and important. 
I followed him in graduate school at the University of 
New Mexico, but did not meet him in person until the 
Texas A&M mammal meetings in 1970, when I had just 
finished at UNM. He was warm and welcoming and 
made me feel a part of his group immediately. I didn’t 
see him again until he hired me to come to the National 
Museum over a year later, while I was on a post-doc 
in Costa Rica. He welcomed my family into his house 
while we looked for housing, and he car-pooled into 
work with me for years, again bringing me into his 
world and making me feel welcome in all circles. I 
witnessed this behavior with everyone who came to 
work for Clyde in those early years of the 1970s. 

Clyde was ambitious and hard-working, and 
worked his way up the ladder in the Fish and Wildlife 
Service, but he always remembered those who worked 
for him and with him. He loved field work and tried 
hard to involve everyone who wanted to participate in 
field activities. He was a good observer, a keen trap¬ 
per, and an excellent preparator, so field work with 
Clyde was always productive and fun. Also, thirst was 
rarely a problem if Clyde was in charge. I think he was 
singularly responsible for the Jos. Schlitz Brewing Co. 
lasting as long as it did. 

Clyde had a favorite watering hole called the 
Crown Bar & Grill, on E Street, just a couple of blocks 
from the Museum. He came to work early every morn¬ 


ing, and put in long days, but he then extended them 
by heading up to the Crown after work. He recruited 
whoever was around for these excursions, from staff 
to visitors. We had to learn to pace ourselves early on, 
as keeping up with the Jones in this case was a difficult 
proposition. Good conversation and beer went hand- 
in-hand for Clyde, and he had an enormous capacity 
for both. Our workforce was diverse in those days, 
and international visitors from every part of the world 
were with us regularly. Clyde treated everyone equally, 
and made sure everyone was involved in whatever was 
going on. 

Although hindsight undoubtedly filters memories 
of 40 years ago, those halcyon days of yore were heady 
ones for all of us who came into contact with Clyde, 
and I believe we were all made better for it. He was the 
best boss I ever had and just had that knack for making 
all of us feel like what we were doing was important 
and worthwhile. He built a terrific team during his 
tenure in Washington, and his presence was missed 
immediately on his departure. Nevertheless, he was 
very good at keeping up with all of us over the years. 
Someone, somewhere in the world, is skinning a mouse 
with a cold beer in reach, and thinking of Clyde and 
what he meant to all of us. 


Richard Manning (p. 67) 


It may not be possible for me to articulate how 
important Clyde Jones (CJ) was in my development 
as a mammologist, and it’s also difficult to talk about 
Clyde without mentioning J Knox Jones, Jr. (JKJ). 
There never is a day, even now, that I don’t think back 
to my training at Texas Tech and the influence CJ and 
JKJ had on my career. 


Clyde Jones and Knox Jones were co-chairs on 
my PhD committee. I was at the NSRL in Knox’s 
office, during the summer of 1985, when I first met 
CJ. Clyde was more the field-guy, while Knox was 
the museum-guy or office-museum-administrator guy. 
Clyde had been gone most of the summer months trav¬ 
eling throughout north-central Texas gathering data on 


233 


234 


Clyde Jones Memorial Volume 


Texas Kangaroo Rat (Dipodomys elator ) distribution 
for Texas Parks and Wildlife Department. 

My first impression of Clyde was that he was a 
friendly, outgoing man with a great sense of humor. 
I remembered Knox later cautioning me don’t 
mistake his casual or informal manner as a lack of in¬ 
terest in your work and academic progress.” True, he 
and Knox, were serious about academics. Both were 
strict concerning field research. It was “drummed into 
me,” early and often, that I needed to take detailed field 
notes, accurately record my observations in the field, 
and most importantly prepare museum-quality mammal 
specimens. I had good instruction. There always was a 
good amount of friendly banter, and advice, while pre¬ 
paring specimens. After each field trip, with or without 
them, all specimens were meticulously examined and 
critiqued, often with a few friendly jibes. Both gave 
suggestions and usually some comment like “Is that all 
you caught?” or “Where are the rest of them?” — usually 
in good humor! I often heard another old adage from 
them: “... when there are a lot rats, get as many as you 
can, and when there are not very many, get them all.” 

As a graduate student these goals were expected: 

(1) to perform at a high academic level in the classroom; 

(2) to show up and work in the mammal collection at 
the Museum (NSRL); and (3) during my “copious and 
significant free time” conduct field work and publish 
the results of that effort. Both Clyde and Knox were 
supportive in my efforts and always available for dis¬ 
cussions and consultation. Their personal libraries 
always were available for my use. 

I took a couple of courses from Clyde (I believe 
a Museum Science class and a couple of seminars and 
a field methods course) and over the years got to know 


him quite well. Only rarely could I get both Clyde and 
Knox out into the field with me at the same time. Those 
trips were some of the most memorable experiences of 
my PhD training. Those trips were filled with continu¬ 
ous stories about their earlier careers; their colleagues 
and friends; their memories of former colleagues; and 
various escapades they had enjoyed with each other. 

Did I mention both Clyde and Knox enjoyed a 
“cold beer” now and then? The more ‘relaxed’ they 
got, the more interesting the stories got. What oral 
history lessons! I wish I had taken a portable tape 
recorder on some of those trips in order to record those 
conversations. 

After I graduated I stayed in contact with Clyde 
and did field work with him, usually in far West Texas. 
He had an intrinsic interest in mammals and was en¬ 
thusiastic about sharing his thoughts. 

He always had ideas about potential mammal 
research projects to discuss. More than once he said 
something like “...we need to look into that a little 
more seriously” or “...let’s try to get a few more of 
those (rats/bats) so we can try and figure out what’s 
going on with them.” 

I knew Clyde as an “academic” father and mentor; 
as an “uncle-like” figure, and finally as a close friend. 
One more thing I’ll always remember him saying — “be 
sure to have fun!” 


Pf 


Mike Bogan (p. 97 and p. 215) 


When I reflect on my time spent with Clyde, 
it seems almost impossible that we were together as 
much as we were and how the times all seem so good. 
Although I’d known about Clyde since 1964, when 
my graduate career in mammalogy started under Gene 
Fleharty at Fort Hays State, I didn’t meet him until 1971 


when I was enrolled at the University of New Mexico 
(UNM). My first field trip with him was in 1974, my 
second summer of working for him and Don Wilson 
at the National Museum of Natural History (USNM). 
My last field outing (but no real work!) was in mid- 
June of 2013 with several friends and colleagues. This 


Encomia and Reflections 


235 


marked over 40 years during which I spent some time 
with Clyde. And I think that much of what I am, and 
have, today is a direct consequence of having worked 
for and with him. Clyde, his research philosophy, the 
opportunities he helped provide, and the organization 
he built were very influential in my life. It was at the 
request of Clyde and Don Wilson that I applied for a 
position with the Fish and Wildlife Service. And later, 
after Clyde moved to Denver, I took advantage of an 
offer from him and moved to Colorado as well. After 
he moved to Texas we stayed in touch, often attending 
meetings together or working together in the field. 

Clyde made many contributions to our under¬ 
standing of southwestern mammals, especially bats, 
while he was a graduate student at UNM. These con¬ 
tributions were a great help to those of us that followed 
him and who were able to use the specimens, field notes, 
as well as publications that he left for us. The rigor 
of his work is shown by the fact that today’s younger 
scientists still commonly cite many of these studies. 

I’ve always been impressed with his work in Rio 
Muni, Africa, on primates and other mammals. The 
work was funded by grants and lasted about three years, 
but it resulted in several of his favorite publications 
and demonstrates Clyde’s abilities and willingness to 
work, as we say now, “outside the box.” Rather than 
“playing it safe” as a beginning Assistant Professor 
in a university in the U.S., I think he saw this as an 
opportunity to conduct research on a new group of 
mammals in a foreign setting and with the opportunity 
to interact with several experts, such as Louis Leakey. 
This work predates my time with him, but I’ve always 
been impressed that he did it. It’s tangible evidence of 
his interest in other mammals, his interest in working 
overseas, and his ability to work constructively with 
others. 

His experience in Rio Muni was valuable to him 
when he went to work for FWS in 1970. His ability 
to interact successfully with a wide array of people 
and agencies in seeking support for FWS research, 
especially on endangered species and marine mam¬ 
mals, was impressive. He also was very successful 
as a research administrator for FWS. That is, up until 
the FWS changed his job and tried to return him to 
DC and he quit. When he quit FWS, many of us were 


very disheartened, probably partly because we knew 
we would have to work harder because Clyde wouldn’t 
be there to do it for us! 

Not surprisingly, as we say, Clyde landed on his 
feet and probably ended up where he always belonged: 
in academia, at Texas Tech University. He started as 
the Director of The Museum, later Chair of the Depart¬ 
ment of Museum Science, and then Professor in the 
Department of Biological Sciences for over 20 years. 
He’s had over 20 graduate students, many of whom I 
know and have spent time with in the field. Clyde has 
acknowledged that without his graduate students he 
would feel that he would have failed at Tech. 

And, the story is incomplete without reflecting on 
the time I spent with him in the field; and now I realize 
that our time together, in many ways, was a gift. We 
were happy to spend the day, skinning trays in our laps, 
preparing specimens that we referred to as “things of 
beauty and joys forever.” I think we envisioned the 
specimens residing in their trays in museums for de¬ 
cades if not centuries-and being of great use to other 
scientists. I don’t know how many hours I’ve spent in 
camp with him doing this. Thousands of hours I am 
sure. And at sites in New Mexico like Willow Creek in 
the Gila National Forest or Cloverdale in the bootheel, 
or Tiburon Island, or Baja California Sur, as well as 
federal lands in Nebraska, Colorado, Utah, Arizona, 
and elsewhere. 

Our last fieldwork together was in the Chinati 
Mountains in far southwestern Texas. We made sev¬ 
eral trips to different sites there and had really magical 
times. It was a nice mix of older and younger scientists, 
all committed to reaching a better understanding of the 
mammals of the Chinatis. And there was the occasional 
sip of “top-shelf ’ bourbon or rye that we brought along 
for after-hours relaxation. 

In the summer of 2013, several of us that had 
worked with Clyde in one capacity or another, got 
together with him and Mary Ann at Big Bend Ranch 
State Park for a few days. Mark Lockwood put this 
trip together, although I don’t think he had anything to 
do with the golf-ball sized hail that did thousands of 
dollars of damage to our trucks! I had recently gotten 
a knee replacement and was trying to do a little hik- 


236 


Clyde Jones Memorial Volume 


ing to strengthen it. Clyde wasn’t hiking at that time 
but twice he came out in his truck after I’d been gone 
awhile to check on me and offer a ride back to camp. 
That was typical of CJ and I appreciated it. 

“Adios CJ-we had a good one.” 


David Schmidly (p. 127) 


I first met Clyde Jones in 1969 while attending the 
annual meeting of the American Society of Mammalo- 
gists. Over the next 46 years I had the great pleasure to 
get to know him as a friend, companion on field trips, 
and as a faculty and administrative colleague. 

Clyde’s life story is fascinating and we are fortu¬ 
nate that it was recorded in his own words. In a paper 
entitled “You Have to Catch Them First,” published in 
the book “Going Afield“(Carleton Phillips and Clyde 
Jones eds., 2005, Museum of Texas Tech University, 
289 pp), he chronicled his life from growing up as a 
boy on a small cattle farm in Nebraska, where he was 
raised by his mother and aunts and uncles; to graduate 
work in mammalogy at the University of New Mexico; 
to an adventure studying African mammals; to an 
accomplished career as a government scientist and 
administrator; and finally as a successful academic 
scientist and administrator. As Clyde put it himself, 
“my personal life in general and my professional career 
in particular have been quite rewarding to me.” 

There are many words I could use to describe 
Clyde—honest, decent, fair-dealing, commonsensical, 
great sense of humor and optimism—but I will always 
remember him for his basic kindness. My wife, Janet, 
and I vividly recall the time he met Janet’s mother, when 
she was an elderly woman in her 90s. He went out of 
his way to talk with her and to make her feel special. 
Afterwards, Mrs. Knox commented to us—what a kind 
and decent man!! And, that is the way Clyde was—a 
gentle person who was kind and helpful. 

At the same time, Clyde lived life on his terms. 
Perhaps it was his growing up on a farm and being close 


to the land. He lived life to the fullest and did it his 
way never compromising his fundamental values. So, 
when James Watt took over as Secretary of the Interior 
and tried to change the philosophy of managing the 
nation’s natural resources, Clyde wouldn’t have it. He 
walked away from his government career rather than 
compromise his values and scientific underpinning. 

Clyde was of the old guard of naturalists—those 
that believed that field work and collecting was es¬ 
sential to natural history. We are losing a generation 
of these field biologists and they are not likely to be 
replaced. Clyde will be missed but remembered for 
his many accomplishments and for living life his way. 

Waldo McAtee, who spent five decades work¬ 
ing for the U. S. Biological Survey, said this about 
the death of valued colleagues, “Merely to recall all 
of these departed comrades is enough to break one’s 
heart, and no cry of woe, however deep, can assuage 
the feeling of their loss.” I can think of no better way 
to express my feelings about my friend and colleague, 
Dr. Clyde Jones. 

Finally, I along with my co-authors, John Karges 
and Robert Dean, feel a need to apologize to Clyde for 
having published a paper in his honor in which we did 
not collect any specimens!! 

S)J 


Encomia and Reflections 


237 


Frank Yancey (p. 147) 


In 1994, Clyde Jones presented to me an auto¬ 
graphed copy of Handbook of the Mammals of the 
South-Central States , a recently released book of which 
Clyde was a co-author. The inscription that precedes 
his signature reads “TO FRANKLIN DELANO 
YANCEY, II, STUDENT-COLLEAGUE-FRIEND.” 
From that point until his death, I was very fortunate to 
know Clyde on all three levels. 

With his reputation and high stature as one of 
the world’s premier mammalogists, clearly Clyde could 
have chosen someone much smarter than me as his 
student. But for reasons I don’t quite gather, Clyde 
agreed to serve as my PhD advisor. As one might 
guess, Clyde was very effective in distributing his 
knowledge of mammals. But what made Clyde such 
an exceptional mentor was his brilliant knowledge of 
the history of mammalogy and his close association 
with so many renowned field biologists. Moreover, 
his willingness to share this knowledge and introduce 
his students to so many of these “legends” was what 
made Clyde truly unique as an advisor. 

Field trips, of course, were a focal point in the 
curriculum of a Clyde Jones student. I recall my very 
first field trip with Clyde to Big Bend National Park, 
at which time I was introduced to the Clyde Jones 
philosophy of problem solving. I had just captured 
a woodrat, and as I was removing it from the trap, 
it clamped down on my hand in the soft tissue area 
between the thumb and index finger, almost penetrat¬ 
ing entirely through with its large incisors. I scurried 
over to Clyde and asked him to help me pry this thing 
off my hand. Clyde responded with a “no problem,” 
and then grasped the rat firmly and, with a sudden and 
direct jerk, ripped the animal from my hand, appar¬ 
ently forgetting the “pry” part of my plea for help. He 
then calmly stated “problem solved... nothing that a 
little duct tape won’t fix.” That was the first of many 


memorable field trips with Clyde, each with its own set 
of unique and humorous stories. 

It wasn’t until after I read the dedication on the 
book that I gave the term colleague much thought. But 
that really is how Clyde treated his students. Students 
were included in all of Clyde’s professional activities. 
We were not only included in Clyde’s research pub¬ 
lications, but, upon Clyde’s insistence, were always 
listed first on the author line (whether deserved or not). 
Clyde’s rationale for this policy was that he was “... not 
the one trying to build a resume.” For nearly 20 years 
after completing my degree under his guidance, I had 
the privilege to continue working on various research 
projects with Clyde, completing a final jointly-authored 
manuscript three days prior to his death. 

Being a student and colleague of Clyde was 
an honor and a privilege, but being a close friend of 
Clyde is what I cherish most. Getting to know Clyde 
on a personal level made for the time of my life. His 
kindness and generosity were beyond what I have 
ever experienced in a friend. The countless field 
trips, skinning sessions, dinner outings, and friendly 
gatherings with an “occasional beer” will live on in 
lore. “Saving rats on the pinning board,” Natural 
Light, 20 minute naps, Yucatecas, navy windbreakers, 
Birkenstocks, Conference Cafe, Whiskey Club, etc. 
bring back the best of memories. Clyde: You, as you 
often referred to a well-prepared specimen, are a thing 
of beauty and a veritable joy forever. Thank you for 
being the best possible mentor-colleague-friend. 


238 


Clyde Jones Memorial Volume 


Art Harris (p. 157) 


As part of the research for The Mammals of New 
Mexico , Clyde and I toured western U.S. mammal col¬ 
lections to document their holdings of New Mexican 
specimens. Clyde was the designated driver, and he 
also acted as gopher in the collections as I, right wrist 
in a cast, spoke into a wire recorder. 

Both of us were in awe of the collections and the 
reigning mammal curators, but it was not all business. 
Clyde had relatives (in-laws, as I recall) in California 
which we were to visit, and one of my clearest memo¬ 
ries is of the elaborate preparation he insisted we make 
before ringing the doorbell. It took careful coaching by 


Clyde and several tries before he concluded we were 
suitably positioned to announce our presence and be 
greeted by his relative—each of us with hand extended, 
carefully sculptured to the exact fit of an expected can 
of brew. Although a serious researcher, he also knew 
how to party. 


Mark Lockwood (p. 177) 


I first met Dr. Clyde Jones in 1994 when he, 
along with Frank Yancey and Mary Ann, came to the 
Texas Parks and Wildlife Department headquarters to 
meet with David Riskind. Frank was conducting his 
dissertation research at Big Bend Ranch State Park 
and the meeting concentrated on incorporating GIS 
into the work. I was familiar with Clyde because he 
has served on a thesis defense committee for a fellow 
graduate student at Sul Ross State University. At the 
time I was investigating the small mammal remains 
from Barn Owl pellets from a grain storage bin on 
my family’s farm in Crosby County but was not con¬ 
fident in my ability to correctly identify the skulls. I 
approached Clyde to ask if he would be interested in 
the project and before I could even finish asking him 
he simply stated “Yes.” My next question was how 
many pellets would make for an adequate sample and 
he responded “I want all of them.” Over the next three 
years I would deliver at least one gallon-sized ziplock 
bag full of pellets, until one day in 1998 I realized that 
“all of them” actually had a limit! 

In 2003 Clyde started a baseline survey of the 
small mammals of Chinati Mountains State Natural 
Area in Presidio County. By that time I was working in 
the Texas State Parks regional office in Fort Davis and 
was involved in coordinating such projects in Trans- 
Pecos parks. I accompanied Clyde, Mike Bogen, Tony 
Mollhagan, and a couple of students into the property in 


the spring to start the project. Clyde was very pleased 
to finally have the opportunity to work in the range, 
and I provided the needed information about access to 
the various areas in the SNA and departed. I made a 
second trip down with Clyde in the summer of 2004 
to discuss how the project was progressing and was 
involved for the first time in trapping small mammals. 
I knew very little about rodents and Clyde noted my 
curiosity and suggested that it would be important to 
the project that I have a larger role and thus began my 
education. I had the great privilege of working with 
Clyde through the completion of the project and sharing 
many days at camp with him, Mike, Tony, Frank, Steve 
Kasper, Rick Manning, and Sam Braudt among others. 
In the latter portions of the project among the many 
rituals surrounding putting up specimens was added 
the reasons why I was close to being a mammologist, 
but not quite there. Clyde would recite that I “trapped 
mammals, pinned specimens, kept a mammal as a pet 
(a Perognathusflavus), and all that was left was for me 
to skin and stuff a specimen.” I am still lacking that 
last part. It was my great pleasure to contribute to that 
project and to be one of Clyde’s unofficial students. 


Encomia and Reflections 


239 


Ken Geluso (p. 183) 


Unlike most contributors to this volume, I never 
had the pleasure of being in the field with Clyde; how¬ 
ever, at meetings and on other occasions, I enjoyed 
Clyde’s friendship and our conversations about bats, 
rats, and other things. And, I always appreciated the 
sound advice and encouragement that he gave me 
through the years. It was Clyde’s study of bats in 
southwestern New Mexico (1966, The American Mid¬ 
land Naturalist 76:522-528) that inspired me to initiate 
a 34-year study of bats in the San Mateo Mountains 


of New Mexico. For many years, Clyde’s greeting to 
me would be, “So Ken, how’s that Bear Trap study 
going?” I thanked Clyde once again soon after the 
study was published in 2012 (Journal of Mammalogy 
93:161-169). 


Robert Bradley (p. 201) 


I first met Dr. Clyde Jones in February 1984. I 
was a young MS student from Texas A&M University 
and was attending the annual meetings of the Texas 
Society of Mammalogists at the Texas Tech University 
Campus in Junction, Texas. Paisely Cato, a PhD student 
at A&M, was being co-chaired by Dr. Jones and she 
introduced the two of us. As I remember, Clyde and I 
were part of a group of graduate students and faculty 
who spent the first night of the meetings socializing on 
the bank of the South Llano River, which flows through 
the TTU campus. I was anxious to meet the man I heard 
so much about and became enamored by his stories 
of field-work and of his association with the icons in 
mammalogy. About every 30 minutes, Clyde tried to 
convince me that the South Llano River was fairly 
shallow and one could probably successfully drive the 
A&M van across it, if one achieved the appropriate 
speed. A few beers later, I countered that it probably 
could be done, but I needed his local expertise for cal¬ 
culating slope of the riverbank, trajectory of proposed 
entry, necessary speed of vehicle across the floodplain, 
and other factors that only a TTU professor, such as 
he, would know. Eventually, Clyde agreed to be my 
co-pilot in this endeavor and a plan was formulated. 
Fortunately, Paisley retrieved a couple of more Natty 
Lites and confiscated the keys. It is debatable whether 
the plan would ever have been implemented, but I did 
gain a life-long friend that night! 

In 1994,1 returned to TTU as a brand new As¬ 
sistant Professor. Clyde was now an “old fart” (his 
words) in the Department of Biological Sciences and 


Curator of Mammals at the Natural Science Research 
Laboratory. His advice and wisdom on how to survive 
as a young faculty member and handle departmental 
politics was generously given and heeded. I learned to 
value our conversations on departmental and university 
happenings. He always knew what was coming down 
the pike and provided the necessary “heads up” when 
needed. He was a valued resource to the Department of 
Biological Sciences and he acted as the go-to resource 
relative to institutional memory. 

Over time, I co-authored 17 papers with Clyde. 
Most pertained to the Revised Checklist of North 
American Mammals North of Mexico series or were 
distributional records and natural history manuscripts 
from our work in the Trans-Pecos region. My favorite 
manuscript was “Molecular Systematics of Dipodomys 
elator (Rodentia: Heteromyidae)”, in which I drug 
Clyde kicking and screaming into the world of mo¬ 
lecular systematics. Often he would good-naturally 
comment to his contemporaries that I had ruined his 
reputation! 

I have many fond memories of working with 
Clyde in the Davis Mountains (see Photos 12 and 13 
in the “Photographs” section of this volume). From 
his license plate (last three digits - 007) that gave him 
a self-proclaimed announcement—“Name is Jones, 
Clyde Jones, and I have a license to kill - it says so right 
there on my license plate”—to his vast knowledge of 
the Trans-Pecos mammal fauna, I learned a lot and had 
many laughs. My students adored Clyde and cherished 


240 


Clyde Jones Memorial Volume 


the times they spent with him, whether it was in the 
field or through his weekly excursions to my research 
laboratory. How he could ever prepare a Perognathus 
flavus specimen with those catcher’s-mitt hands is still 
a mystery! He “salvaged” many of my specimen-preps 
and turned all of our specimens into THOBs! 

Clyde, I miss you. I hope your traps and nets 
are always full and may you never run low on cotton 
and wire. 


Tony Mollhagen (p. 215 and p. 97) 


Clyde Jones and Gene Fleharty were undergradu¬ 
ates and friends at both Hastings College and later at 
the University of New Mexico. After leaving New 
Mexico, Fleharty took a position at Fort Hays State, 
where I was enrolled. He was my supervisor, first as a 
work study undergraduate student, then eventually as 
his Assistant Curator in the mammal collection. He was 
also the advisor for my master’s thesis. I had heard 
many, many stories about Clyde Jones long before I 
ever met him. 

In 1965, Fleharty took me, Elmer Bimey, and 
John Farney on a collecting trip to New Mexico. One 
of our camps was at a then famous Euderma locality, 
Willow Creek, in the Mogollon Mountains. It was there 
I met Clyde. He drove from Albuquerque and arrived 
in camp after dark. After introductions, he took drink 
orders. In the trunk of his car was a homemade, built-in 
primitive bar with a wooden rack for bottles, a cutting 
board, and a cooler with ice and condiments. He and 
Fleharty just had Black Label Jack; I don’t remember 
what the rest of us took. They told stories and we 
laughed. Clyde spent only the one evening with us. 

In 1967,1 came to Texas Tech as a PhD student of 
Bob Packard. By then Clyde had taken a job at Tulane. 
Before my first class at Tech I was in a conversation 
with a new faculty member in the mailroom, Francis 
Rose. Rose had come from Tulane, so I asked if he 
knew Clyde. He most certainly did and we exchanged 
several stories. Shortly, I learned my TA assignment 
had been changed from freshman Zoology lab to a 
premium assignment, Rose’s Anatomy and Physiology 


lab. Presumably simply knowing Clyde Jones was an 
endorsement. 

For years after our meeting at Willow Creek, I 
had only minimal contact with Clyde. This was true 
even after he came to Tech in 1982. The encounters 
were chiefly at meetings or in the company of a visiting 
friend, Mike Bogan, who had worked with Clyde for 
many years. While my own career had substantially 
deviated from my earlier training, Clyde generously 
asked my opinions as though I was current in all things 
mammalogical and I could possibly contribute to a 
discussion. 

Nevertheless, it was during this period (1989) 
I shared my first author line with Clyde, a result of a 
capture of a range extension of a Harvest Mouse some 
time earlier. A person has to be a certain age, and have 
a certain academic heritage, to appreciate having one’s 
name associated with that of even one of the Jones boys 
(Clyde and J Knox), much less both. Never mind that 
it was a minor pub and my name was an afterthought 
in a string of authors. I had published more significant 
work in other disciplines, but I admit to being a little 
puffed-up over that one. 

My next and last shared authorship with Clyde 
resulted from the only period I actually worked in the 
field with him. This was his Chinati Mountains project, 
published in 2011. I was retired from the faculty in 
Civil Engineering, but still had some skills at collecting 
mammals and prepping specimens, thanks to a long 
association with Bogan. Most commonly the trips to 


Encomia and Reflections 


241 


the Chinatis consisted of small to large crews of current 
or former colleagues and current or former students, 
but sometimes it was just me and Clyde. Whatever 
the case, whenever the serious work was done, the 
evenings usually concluded with Clyde holding court. 
There was laughing, story-telling, and beverages over 
ice; Knob Creek was favored at the time, among other 


distillates. The circumstances were not unlike our first 
meeting over 40 years earlier. 


Jim Goetze (p. 225) 


My family has a fairly long association with Dr. 
Clyde Jones. This association was initiated when Dr. 
Jones first began sampling small mammals, and in 
particular the Texas Kangaroo Rat (Dipodomys elator ), 
on some Goetze family properties in Wichita County, 
Texas. This occurred when Clyde was a faculty mem¬ 
ber at Tulane University. As Dr. Jones told me later, “I 
first met you when you were about knee-high.” 

My next brief encounter with Dr. Jones was 
when I traveled to Lubbock around 1985 to research a 
Master’s Degree in Museum Science from Texas Tech 
University. At that time, Clyde was the Director of 
the Museum of Texas Tech University and intimately 
involved in the Museum Science Program. At this time, 
circumstances intervened and I ended up not pursuing 
that particular pathway. 

I didn’t meet Clyde again until the summer of 
1989, when, once more, I traveled to Texas Tech Uni¬ 
versity to interview for a spot in the Doctoral program 
of the Department of Biological Sciences. My purpose 
was to interview with Clyde and J. Knox Jones concern¬ 
ing their program and my particular research interests. 
I don’t remember too many specific questions from 
that interview, but one, duel question is still crystal 
clear in my mind; and it was this, “Why, in particular, 
do you wish to pursue a PhD with us, and what do you 
intend to do with it, IF you earn one?” Now, I don’t 
remember my exact answers to those two distinguished 
scientists, but recall telling them that I had developed 
my current interest in biological sciences while attend¬ 
ing Midwestern State University and, initially, taking 
courses for a teaching certification. As a result, I had 
become very interested in the discipline (particularly 
in field work, museum specimen preparation and care, 


and curatorial activities in a natural history collection) 
and had decided to obtain a Master’s Degree in Biology 
from MSU and perhaps pursue a doctorate degree, if 
accepted. In fact, my paleontological-based, reworked 
master’s thesis had recently been published in the Texas 
Journal of Science, of which both Clyde and Knox were 
involved in peer-review, editing, and the publication 
process of that journal. I told Clyde and Knox that I 
really enjoyed the field research and publication aspects 
of biological sciences. 

What sticks in my mind about this is that they 
both smiled and allowed that I had given them a ‘good 
answer’ because, if I had said “I want the degree to 
teach at a college or university” then I would not 
have been accepted into their program. Clyde and 
Knox demanded that their graduate students conduct 
independent research, and that this research had to be 
“publication grade.” In addition to their graduate stu¬ 
dents’ own thesis or dissertation research, Clyde and 
Knox also expected their students to participate in other, 
numerous projects that they (Clyde and Knox) initiated. 

I was subsequently accepted into their program, 
and thus began a longer and closer association with 
Clyde. As stated before, with Clyde most of your own 
graduate research was expected to be independent and, 
despite wracking my brain for memories, I can’t think 
of an instance where Clyde was involved in my dis¬ 
sertation fieldwork on the Edwards Plateau of Texas. 

HOWEVER, I was associated with Clyde on sev¬ 
eral other field-based, research projects including the 
mammal and herpetological survey work on the Harte 
Ranch addition of Big Bend National Park, the Lake 
Meredith National Recreation Area mammal research 


242 


Clyde Jones Memorial Volume 


project, the Lake Allan Henry environmental impact 
study for the City of Lubbock, and the non-field-based 
publication of two Mammalian Species accounts and 
a species account for the Smithsonian Book of North 
American Mammals. 

The first thing that I can say is that I thoroughly 
enjoyed working with Clyde on all of these projects 
(field-based or not). It seemed to me that Clyde was 
always happiest when conducting fieldwork and, as 
everyone who knew Clyde Jones can attest, he was a 
consummate, professional field biologist and a “speci¬ 
men preparator” par excellence! 

Also, it was a great deal of fun to be in the field 
with Clyde; where he never seemed to lose his sense 
of humor (rain, shine, hail, cold, or come-what-may) 
and always performed more than his share of the tasks. 
Clyde was almost always the first one up in the morn¬ 
ings, and was usually cooking everyone breakfast (with 
the coffee pot on—although Clyde often was drinking 
some ‘liquid cereal’ because we all knew that, “One 
beer is worth 20 minutes of sleep.”) with the usual 
small smile on his face and a cheerful greeting. We 
would be “out the door” to take down mist nets at the 
break of dawn and “down the road” for the Sherman 
traps or Museum Specials, depending upon the season 
of the year. 

I especially enjoyed Clyde’s system of “assembly 
line” preparation of the day’s catch. Each field party 
member had a job or part in the overall specimen prepa¬ 
ration that the individual was at least pretty good at, 
and, using this system on a daily basis, large numbers 
of specimens could be prepared, pinned, and stored for 
drying in a most time-efficient manner. Clyde often 
ended up as one of the people with the final task of 
pinning out the voucher specimens for drying. This 
was the time of the famous lines from Clyde that all 
of us remember! 

One: “Well... maybe I can save this one on the 
pinning board... ” and Two: “It’s a thing of beauty and 
a joy forever!” 

One collecting trip memory that I can recount 
(as common in fieldwork, others are perhaps best left 
unwritten) occurred during the Harte Ranch survey. I 
was on this particular collecting trip with Clyde and 


Rick Manning. We drove the old, Dodge, university 
van out from our base camp in the early evening looking 
for a likely place to set some Shermans where we had 
not previously trapped. This is the Chihuahuan Desert, 
so all was dry. Therefore, we drove off of the travelled 
road and some distance through a creosote flats area 
until we reached a small arroyo. Upon inspection of 
the area, we all agreed that this was the “sweet spot” 
and we laid down the traps. I don’t remember exactly 
how many traps, but I would guess a minimum of 300 
because there were three of us. At the time, 100 traps 
per mammalogist seemed to be the standard minimum 
number for each set or transect. However, we probably 
set more traps because we were in a ‘new locality. ’ 

Well, we went back to base camp at Mountain 
Lodge and roosted for the evening after a stint of night¬ 
driving road surveys for nocturnal mammals and any 
herpetological specimens we could find. During the 
night, an intense thunderstorm rolled through the area 
and soaked everything down really well. 

The next morning, we then attempted to reach 
our traps in the old, non-four-wheel-drive van and soon 
discovered that we had better give up on that endeavor 
and start walking to the trap lines. Needless to say, 
the traps were some distance away from where we 
abandoned the university van. Now I can’t speak for 
Clyde or Rick Manning’s footwear, but by the time we 
reached the trap lines, my boots had grown about three 
sizes larger and were somewhat heavier than when we 
began our trek. We picked up all of the Sherman traps 
(empties in the wooden trap boxes and ‘full ones’ in 
canvas trap bags) and started slogging out toward the 
van. About half-way back, my boots are now six sizes 
bigger with the mud and muck on them and much, 
much heavier. At this point, I made what I felt to be a 
relevant comment regarding my condition and Clyde 
smiled and laughed. However, the Boss—as we called 
Clyde—never forgot it, and would occasionally remind 
me of it when times got tough! I just told CJ that, “Hey, 
I can hear my heart pounding in both of my ears now! ” 

We made it back and all was well. Clyde would 
always make sure of that with his watchful eye and 
concern for his students, and I hope, if he is watching 
now, I have made him just a bit proud of his “more 
deliberate” graduate student. 


Encomia and Reflections 


243 


So long, Boss, and I hope that where you are at, 
your trap lines and mist nets are as-full-as you want 
them, your shotgun never misses its mark, and you are 
still making “things of beauty and joy forever!” 


Allan Nelson (p. 225) 


I met Clyde Jones for the first time at the Texas 
Society of Mammologist meetings at Junction, Texas. 
We ended up sitting at the same table after the banquet 
tables had been moved to the periphery to make room 
for dancing. We introduced ourselves and he pulled 
out a bottle of bourbon and offered it to add to the cola 
I was sipping. I thanked him, added a slug and settled 
in to watch the dancing. He sat for a few minutes and 
then said, “I really liked your paper in the Museum Oc¬ 
casional Papers series, where you compared the flora 
of the Texas barrier islands.” 

First this struck me as incredibly nice and sec¬ 
ondly, I was amazed he remembered a rather obscure 
botany paper. Next he began to talk with me about 
the paper recounting details and discussing his own 
thoughts regarding the biogeography of plants and 


mammals in Texas. I remember this because it made 
me feel that others, even people outside my field, were 
looking at something that I did. I guess you could say 
it sort of gave me a mid-career boost regarding my 
research. 

As the night continued, Clyde did this same sort 
of thing with numerous individuals that visited the 
table to talk or get bourbon. There is no doubt in my 
mind that few field biologists that work in Texas today 
have the breadth and depth of knowledge that Clyde 
displayed that night. 

^ 0 . 


Larry Choate (p. 225) 


Writing an encomium for Clyde Jones is espe¬ 
cially difficult because the very act of putting words on 
paper acknowledges that he is gone—difficult to believe 
because, in many ways, Clyde had a larger-than-life 
persona. A scientist of international reputation whose 
published works encompassed the breadth of mammal¬ 
ogy, he also was involved in and published in a variety 
of other disciplines. For example, before I became his 
graduate student, he auditioned me by having me ac¬ 
company him for a week on a month-plus-long trip with 
Royal Suttkis and three graduate students who were 
engaged in sampling fishes in the Red River drainage 
in Oklahoma and Texas. What an eye-opener: endless 
seining in daylight and dark, lots of heat and sand, and 
even more cold beer—sometimes not so cold. 


Clyde was unique in so many ways. His pro¬ 
fessional career, as a researcher in equatorial West 
Africa, with the U.S. Fish and Wildlife Service, and 
in academia was inundated with big personalities with 
even larger egos. Clyde Jones had an excellent sense 
of propriety. I never heard him ‘blow his own horn;’ 
he did not have to because only the most dim-of-bulbs 
were unaware of his status and accomplishments. Still, 
it was never his style to brag. Many times I was with 
him in the presence of others holding forth their theses, 
opinions, or learned postulations while he remained 
stoic. If directly questioned, Clyde would either agree 
or diplomatically provide an alternative explanation or 
solution. Rarely would he take someone to task, and 
never without substantial thought and justification. He 


244 


Clyde Jones Memorial Volume 


did, however, enjoy recounting amusing anecdotes, pri¬ 
marily regarding field exercises that did not go exactly 
as planned. To refer to Clyde as “Old School” was truer 
than most realized. He knew and had interacted with 
many of the first modem mammalogists, and with many 
senior scientists from other disciplines. Clyde had been 
in “the field” with many of these folks. 

Clyde had an exceptional talent for leadership: 
getting others to WANT to do what he wished them to 
do. And, his outstanding interpersonal communication 
ability spanned the human spectrum. Whether he was 
speaking in Spanish to a construction crew, working 
a cocktail party fundraiser, or interacting with peers 
at a scientific meeting, Clyde got things done and, 
usually, everyone left happy. Although he was adept 
at these exercises in leadership, Clyde seemed not to 
enjoy many of these people-interactions as well as his 
solitude. Many of Clyde’s value systems were rooted 
in the past, from his Nebraska ranching heritage. 

But, his range of interests was more varied than 
most would imagine and always included a forward- 
thinking element. While in his company, I often was 
somewhat taken aback when he received an “old friend” 
acknowledgement from people outside of academia in 
everyday life. People such as ranchers at the end of a 
rutted road, florists, painters and sculptors, poets, gar¬ 
deners, hardware store clerks, band members at a coun¬ 
try music concert or dance, and many others seemed to 


genuinely enjoy recognizing Clyde’s friendship. Even 
his knowledge in his chosen discipline of mammalogy 
was extremely varied; for example, among many taxa, 
Clyde was an expert on Primates and marine mammals, 
especially Cetaceans. He loved flowing water and was 
a river boatman of some experience and skill. 

Clyde always had a strong interest in the arts, 
and thought this would be a much better world if our 
public education system did a better job of introducing 
everyone to the arts. In many ways, Clyde Jones was 
a Renaissance man; however, he did have more than a 
few quirks. He used to keep an ornate box turtle as a 
free-range, backyard pet, and he went through a period 
of not answering a telephone. Pavlov notwithstanding, 
it is disconcerting to attempt to carry on a conversation 
while ignoring a ringing telephone. For a few years, 
he also would only drink Schlitz Beer. Schlitz! (and 
then he changed to Miller Lite. Lite!) If Clyde thought 
you needed to sort an answer out for yourself, he would 
not directly answer a question; rather, he would assign 
you an oblique task which (in his mind at least) was 
supposed to lead you to the correct answer. 

I miss him. 


Lou Densmore 


What can I say about Clyde Jones? He was a 
friend, a colleague, a confidant, a mentor and an advisor. 
Not being a mammologist, I do not have stories about 
skinning ‘rats’ or traveling to Africa with Clyde that 
so many folks do. To be truthful, I really did not get 
to know him well until about 15 years ago. Everyone 
knew Clyde by reputation, as he was one of the truly 
iconic mammalogists of the last half of the 20 th century 
(and Texas Tech had three of them, Clyde, Knox Jones 
and Robert Baker). However, until about 2000, our 
relationship was friendly but a bit distant. 

Maybe that is because I had several interactions 
with Clyde early on, some of which were not that 


cordial. One of our first ‘encounters’ was in the early 
90’s. My then fiance (and now wife of 21 years), Erika, 
had driven my vehicle over to the University for some 
long forgotten reason. We had assigned parking spaces 
in Biological Sciences in those days and someone had 
taken my space. Erika was only going to be there for 
about 15 minutes, so she just parked in the space next 
to mine, which happened to be Clyde’s. Dr. Jones 
showed up about 5 minutes later, and needless to say, 
he was not very happy about it. The next time he saw 
me in the hall (which just happened to be in front of 
the faculty mailboxes outside the main office), I got 
‘dressed down’ for about 10 minutes. When Clyde was 
upset he rarely minced words.and true to form, he 


Encomia and Reflections 


245 


did not on that morning. Needless to say, neither Erika 
nor I ever parked in Clyde’s parking place again. In 
later years, we all laughed about it, as Clyde and Mary 
Ann came to be some of our closest friends. 

Clyde had an incredibly keen sense of humor, 
sometimes caustic, but at the same time very dry. There 
were times when he would say something to you and 
after the conversation was over and you were walking 
away, the realization hit that you had just been verbally 
eviscerated and were now ‘tripping over your own 
entrails’ — and at the same time you were laughing 
about it. I remember several instances when his short 
quips, stated totally off the cuff and at the perfect time, 
would just about bring me (and others if they heard 
them) to tears. One example occurred during a faculty 
meeting that dealt with bringing into our department 
an entire center of researchers recruited to Texas Tech 
from another university. As per usual, we had consider¬ 
able discussion about the matter, with both pro and con 
opinions being voiced. After one person’s rather nega¬ 
tive and frankly ludicrous comment, Clyde (who was 
sitting next to me) leaned over and stated something 
to the effect: “isn’t it terrible what chlorine bleach can 
do to the human brain” (let’s just say the person was 
not a true blonde and leave it at that). I literally had 
to “chew” on my arm to keep from laughing out loud. 
I knew at that moment we would be friends forever. 

After Clyde retired, I took over his position as 
Associate Chair. He was still a regular visitor to the 
department and that is when I truly started to benefit 
from his counsel and advice. His impact on me was 
even greater when I became Chair in 2009. Clyde had 
truly just about ‘seen it all’ in his various roles with the 
federal government, in museum administration, and 
in the department. He had the remarkable ability to 
recognize and ferret out the difference between admin¬ 
istrative BS and legitimate (and important) issues. To 
a new chairman sometimes struggling to know where 
to devote his time and energy, Clyde’s advice on such 
matters was critical. 


Probably my favorite moments spent with Clyde 
were during his last years when he and Mary Ann 
would invite Erika and me to come to the Frazier 
Alumni Pavilion on football Saturdays to sit at their 
table in a cordoned off area, consume adult beverages 
(it was Clyde that first introduced me to Knob Creek 
bourbon), and watch the Texas Tech and other football 
games. Clyde has his own “uniform” and everybody 
who knew him can relate to it. He preferred to wear a 
cream colored to tan TravelSmith “Great Escape” style 
short-sleeved shirt (see Photo 26 in “Photographs” 
section of this book) that was never tucked in. His 
pants were normally blue jeans (or sometimes shorts 
for August or early September games), but his piece 
de resistance was that pair of Birkenstock sandals that 
were worn (obviously with no socks) whether it was 
95 degrees or 35 degrees outside. I do not think that 
anyone knew the actual age of those shoes, but they 
were definitely ‘broken in’. If it got bitterly cold, one 
would see him in the same outfit, but also wearing 
what Mary Ann describes as ‘very well worn’ blue or 
black windbreaker. 

Clyde and Mary Ann were clearly adored by the 
Frazier staff and typically allowed to come in a bit early 
and sit at their centrally located table. And it was obvi¬ 
ous that those feelings were reciprocated. They knew 
everyone by first name, from the janitors, to the police 
officers, bartenders and administrators, and there was 
genuine affection displayed by these people for ‘Dr. and 
Mrs. Jones’. During the most recent (2015) football 
season, after his passing, people regularly came up to 
talk to Mary Ann, often tearfully conveying their sin¬ 
cere condolences and relating their own ‘Clyde stories.’ 

Clyde Jones was a man that could be deadly se¬ 
rious in one breath and hilariously funny in the next. 
Honest to a fault, the better you knew him the more 
you began to understand and appreciate his perspective 
and approach. It is perfectly clear to me why he was 
so beloved and is so greatly missed by so many people. 


Gene Fleharty 

It was in August 1953 that I first met Clyde. We became first string but subbed a great deal and did an 
were both on the Hastings College football team— excellent job at guard. In 1954 we were both on the 
Clyde, a freshman, and I, a sophomore. Clyde never undefeated team that played in the Mineral Water Bowl. 


246 


Clyde Jones Memorial Volume 


I graduated in 1956 and became a student of Dr. J. 
S. Findley at the University of New Mexico. The next 
year I tried in vain to get Clyde to apply to UNM. He 
had accepted an assistantship in the physical therapy 
program at Southern Illinois, as I recall. I called Clyde 
and begged for him to send in an application to UNM. 

I didn’t think he would apply, but in a week or 
so Dr. Findley called me into his office and asked if I 
knew a Clyde Jones. I assured him I did and gave him 
my best recommendation. Findley wanted to know 
what the statement “academic suspension” meant on 
his transcript. I told him that Clyde was found drink¬ 
ing beer on campus which was a definite “no no” at the 


Steve 

My experiences with Clyde Jones are bimodal, 
first as an educational advisor and teacher to a student 
and, years later, as a friend and confidant. I first met 
Clyde Jones while I was a graduate student at Mid¬ 
western State University (MWSU). He and several 
of his students were examining mammal specimens in 
the MWSU collection, then under the supervision of 
Dr. Walter W. Dalquest. My first impression was that 
Clyde’s students were devoted to him. Years later, after 
I arrived at Texas Tech University (TTU) in the fall of 
1991, Clyde spoke to incoming graduate students at 
the beginning of the semester. His witty and insightful 
comments were comforting, but I was impressed by his 
quiet confidence. 

My first fieldwork experiences with Clyde were 
during a summer field mammalogy course being 
team-taught by Drs. J. Knox Jones and Clyde Jones; 
we had several field trips during this course, including 
the upper panhandle of Texas, the City of Lubbock’s 
reservoir mitigation lands east of Justiceburg, Texas 
(which, ironically, is where I presently work), and the 
Harte Ranch mammalian survey project for Big Bend 
National Park. Clyde was always quietly in charge 
because everyone wanted to make sure that he was 
pleased. For me that sentiment never stopped. Dur¬ 
ing the years at TTU, I was privileged to have been his 


Presbyterian school. Findley sort of overlooked that 
and eventually Clyde came to UNM. 

In August of 1957 he immediately went with me 
to collect chipmunks in the Guadalupe Mountains in 
southeastern New Mexico. I had to tell him what a 
chipmunk was, and Clyde collected his first specimen. 
Over the next few years we spent much time together 
taking numerous trips to mountain ranges in New 
Mexico as Dr. Findley hired us as field assistants on a 
NSF grant to study the mammals therein. 

Many happy hours were spent with Clyde. We 
both had experiences that are probably best told after 
a few drinks. 

Kasper 

teaching assistant for the labs of two courses he taught 
(Vertebrate Structure and Development and Natural 
History of the Vertebrates). 

We lost touch after I left TTU and went into the 
private sector in Lubbock. Then, around 2005, Clyde 
showed up at my place of business regarding a gift for 
his lovely wife Mary Ann. After some remembrances 
of field work, he stated “Why don’t you come visit 
me?” Over the following years, I went to Clyde and 
Mary Ann’s home countless times to sip some refresh¬ 
ments and just discuss life, politics, TTU and Nebraska 
football, and of course biology. Clyde subsequently 
invited me to accompany him on field projects to the 
Chinati Mountains State Natural Area and Big Bend 
Ranch State Park. His legacy persists to this day with 
the continuing research on both of these projects. 

Clyde’s breadth of overall knowledge and the 
mammalian literature was remarkable and never left 
him. I was always impressed with this facet of Dr. 
Jones during our mammalian discussions. Our friendly 
debates at his home were always entertaining, often 
educational, and therapeutic for the both of us. Clyde 
was a kind, thoughtful man who would have found his 
way in anything he decided to undertake. We are all 
fortunate that he chose the course that he did. 


Encomia and Reflections 


247 


David Riskind 


All my professional life, starting in the mid-60s, 
I had heard of Dr. Jones, the eminent mammalogist. 
Through him, I educated myself of the mammalian di¬ 
versity and distribution of mammals in Texas. Dr. Jones 
helped me greatly with some publication difficulties I 
was having with the Texas Journal of Science. I regu¬ 
larly used the Checklist of North American Mammals 
North of Mexico, and the regular revisions, published 
through the Occasional Papers of the Museum at Texas 
Tech University. In about 1970-ish, I snapped to the 
fact that there were two of them (Dr. Clyde Jones and 
Dr. J Knox Jones)! And, it was my good fortune (and 
the good fortune of the Natural Resources Program, 
Texas State Parks) to finally work directly with Clyde 
Jones. I met him at a ‘Small Mammal Workshop 5 1 had 
organized with Dave Schmidly at the newly acquired 
300,000 acre Big Bend Ranch State Park (BBRSP). 

Clyde, his students, and his colleagues were the 
core of the field and lab crew running us through the 
paces of field collection methods and the specimen 
preparation techniques for documenting small mammal 
diversity and distribution for the newly acquired, and 
largest, state park. The workshop formed the basis for a 
relationship with El Professor, and the Natural Science 
Research Laboratory (NSRL) of the Museum, Texas 
Tech University. Clyde became the nexus for a long¬ 
term project to establish and conduct baseline small 
mammal investigation on Texas State Parklands. The 
core of the effort would be a series of studies leading to 
MS and PhD degrees. The Natural Resources Program 
provided the funding base; TTU, Dr. Jones, the Biol¬ 
ogy Department, and the NSRL provided the graduate 
students, curation, and academic spine. It didn’t hurt 
our effort that Dr. Schmidly became Vice President for 
Research, and then President of TTU, about this time. 

In addition to Dr. Jones and his cadre of faithful, 
enthusiastic, and talented students, we also got Clyde’s 
long-term colleagues, former students, and friends— 
among them Richard Manning, Mike Bogan, Tony 
Mollhagen, Jim Goetze, Larry Choate, Sam Braudt, 
and of course Clyde’s wife, Mary Ann. 

Past forward and we have as a result long-term 
plots established at Big Bend Ranch State Park, Chinati 


Mountains State Natural Area, Davis Mountains State 
Park and TNC’s Davis Mountains Preserve, Caprock 
Canyons State Park, Lost Maples State Natural Area, 
among others. Lranklin D. Yancey, II, currently is 
engaged in revisiting his dissertation research and, as 
an honor to Clyde, continuing our investigations of 
small terrestrial and volant mammals of the Chinati 
Mountains SNA and Big Bend Ranch. 

Along the way we had some absolutely terrific 
field days (and nights) and some memorable moments 
that will be fodder for legend and lore. The 4X4 
adventure trip to conduct field investigations along 
Terneros Creek, BBRSP, Presidio County, stands out. 
Sitting around the campfire enjoying friends and the 
evening sky, Joel Brant hops up, grabs his shotgun, 
and “ON THE PLY” collects a new record! Clyde 
dutifully demonstrates to Jana Higginbotham the fine 
art of specimen preparation. 

San Antonio Cabin at Chinati Mountains SNA 
was a favorite of Clyde’s. It provided an outstanding 
field lab/camp and came with its own population of 
Pallid Bats. The composition and seasonality of that 
night roost became the subject of detailed analysis. 

I had for more than four decades studied the flora 
and vegetation of northeastern Mexico and especially 
the Sierra Del Carmen just south of the Chisos Moun¬ 
tains. Clyde had always wanted to go and, finally, in 
2007 was hosted by our good friends Bonnie and Billy 
Pat McKinney. I guided him and Mary Ann there. 

The trip wasn’t especially long but because the 
terrain was unfamiliar, Clyde was anxious to arrive. 
A blown and ruined tire, along the way, didn’t help. 
We arrived at Pilares, and were greeted by Bonnie. I 
recall that about 2.5 minutes after we arrived Clyde 
said “Where are the specimens?” Recall the only small 
mammal records in the literature were Rollin Baker’s 
study published in 1956. Clyde was especially inter¬ 
ested in the Coahuila Mole and Miller’s Shrew. Bonnie 
had recent specimens of both, and Clyde wasted ‘NO’ 
time in examining them. He was giddy, and then Bon¬ 
nie pulled out the Leptonycterisl 


248 


Clyde Jones Memorial Volume 


My conversations with Clyde, even in the 
evenings were brief, as I recall. He communicated 
frequently by expression. I especially enjoyed his eye- 
rolls (these told you exactly what he thought). As a born 
and reared Plainsman, he usually was guarded in his 
opinions—but the eye-rolls told the story. For actual 
news I relied on Mary Ann; and, of course, she told all. 

I never could get much out of Clyde via e-mail. 
A greeting, one sentence, and closing were about it. 

I attach one of his lengthy “missives”: 


Wed 11/7/2012 10:16 AM 

cjmajQnes@aol.com 

To David Riskind 

0 You replied to this message on 11/7/2012 2:49 PM. 

Dear David: 

Nice job with regard to Bonnie's book 
Best to you, 

Clyde 

I miss them, short as they were! 

I am hopeful that decades from now, someone 
will revisit the work spearheaded by Clyde and his col¬ 
leagues and gauge how our parkscapes have fared - for 
better, or worse. They will be able to do so because 
of his efforts. 


Fred Stangl 


I was a second-year graduate student at Texas 
Tech under Robert Baker, and life couldn’t have been 
better for a young rat-stuffer. In addition to my major 
advisor and the students who shared the lab with me, 
the Biology Department and Museum were staffed with 
the likes of mammalogists J. Knox Jones, Jr., Dilford 
Carter, Mike Willig, and even a young Ron Chesser. 
How could things get better? 

Then, Clyde Jones came into town to interview 
for the directorship of The Museum. His presentation 
on mammals of the Grand Canyon’s Colorado River 


was promising. Even more promising was what I saw 
shortly after he assumed the job: feeling the need to 
take off for a weekend, this museum administrator 
headed off into the northern Panhandle and came back 
with pinning trays full of mammals. 

Among all those rats and mice, I remember 
stuffed examples of a coyote and striped skunk. This 
kindred spirit came along a little too late to serve on 
my graduate committee, but he was a good friend and 
valued mentor throughout my career and continuing 
on until after my own retirement. 


Richard Stevens 


I met Clyde in the Fall of 1990. I took Mam¬ 
malogy from him and it changed my life. The next 
semester I took Vertebrate Structure and Development 
from him. Perhaps halfway through the semester he 
pulled me aside after class and said that he had a gradu¬ 
ate student conducting a biodiversity survey of a new 
extension to Big Bend National Park (Harte Ranch) 
and asked if I would go on the next trip and help him 
out. The first night we placed a mist net over a metal 


overflow tank right outside the house at Harte Ranch 
and caught among other things, a Townsend’s long¬ 
eared bat (Corynorhinus townsendii ). That moment too 
changed my life for I have worked on bats ever since. 
Clyde and J. Knox Jones Jr. (Los Cojones) nominated 
me to the American Society of Mammalogists back in 
the day when such a formality was necessary. This 
remains my “home society” and has had a huge place 
in my life for the last 25 years. Clyde was responsible 


Encomia and Reflections 


249 


for much of the mammologist I am today. He taught 
me the nuances of trapping and preparing, though you 
would not know it by comparing my specimens to his. 
He also taught me how to drink Natural Light... for 
breakfast, that when you are really tired and hot in the 
field a beer is equal to a 15 minute nap, and perhaps 
most importantly, not to try to keep up with him. Clyde 
was one of a kind. He was also one of a cadre of tra¬ 
ditional mammalogists from another time; the golden 


age of Mammalogy. My experiences with Clyde were 
some of the most important to the development of my 
career. Through those interactions, the stories, and the 
pedagogy on how to study mammals, I was connected 
with this golden age. My time spent with Clyde Jones 
was a true honor though not sufficiently appreciated at 
the time. It’s not that I was unappreciative. It is that I 
did not have the capacity to fully appreciate the giant 
I was amongst and how truly fortunate I was. 


Ernie Valdez 


It was the early 1990s and I was just a neophyte 
to mammalogy when I went on my first major field ex¬ 
pedition to the Sandhills of Valentine, Nebraska, to trap 
small mammals. That is where I met some “giants” in 
the field of mammalogy, with one of them being Clyde 
Jones, but I didn’t really know much about them or their 
greatness at the time. Being young and impressionable, 
I was like a sponge in absorbing knowledge from Clyde 
and others while in the field. It also helped that Clyde 
could see my desire for mammalogy and willingness 
to learn as much as I could and he was happy to pass 
on his knowledge. 

We would spend mornings picking up traps that 
were set the night before and return to camp to prepare 
them as museum specimens. During our morning prep 
of rodents that included Peromyscus, Dipodomys, 
Onychomys, Microtus, and more, we would set up our 
assembly line and prepare specimens together. Willie 
and Family Live would be playing in the background, 
then you’d hear the crack of a beer, and fur would be 
flying, so to speak. During this time is when I would 
hear the first of many sayings by Clyde such as, “skin¬ 
ning mice gives me gas” after a low burp. Other sayings 
included comments about rodents that I had never seen 
before in my life, such as seeing Onychomys for the 
first time. In particular, we would skin them out and 
then collect reproductive data from them. As many of 
you know, male Onychomys can have large testes when 
they are in breeding condition, and during our prepping 
Clyde would make the sympathetic comment towards 
them by saying, “Did ya see the size of nuts on that 
guy? Oh my God,” as he would cup his hands out as 
if holding two grapefruits. These sayings would make 
me laugh and engrave a long-lasting, great memory 


into my mind, but also helped me remember the genus 
Onychomys by association. 

Clyde also had a major impression on me when 
it came to prepping rodents. Although he was skilled 
at skinning small mammals, he was the grand master 
of stuffing and pinning rodents. I could see that he 
would take pride using pins to hold back the smallest 
hair that was out of place and was keen on giving the 
rodents square butts, while telling folks that he could 
save our specimens on the board. He set the bar high for 
me when it came to the quality of preparing a museum 
specimen; a dying art that has since taken second or 
even last ranking to preparing fluid/alcohol specimens, 
taking only fur clips and biopsies, or collecting genetic 
material first while letting the skins dry out or become 
greasy from melting fat. Fortunately, he instilled the 
importance of having a quality voucher specimen that 
can be used in perpetuity and did so by being frank 
about his critiques related to my museum preparation 
of specimens. 

One of my fondest memories of his frankness 
was when he had given me my first Neotoma to s kin 
and stuff. This was a large adult and should have 
been easy to prepare but after spending more time on 
it than I should, I finally had a rodent that was stuffed 
and wired. In my proud but humble moment, I asked 
Clyde what he thought of my work before I sewed it 
closed. His response was, “Jesus Christ! This looks 
like shit! I give you a piece of cake; all I wanted was 
a little cherry on top. Go ahead and do it again, ok?” 
I smiled at him and he back at me, knowing that this 
was honest but tough love. Since then, I have worked 
over the decades perfecting my prepping skills, as 


250 


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well as my impersonation of his voice and this saying. 
I often share this story with young students learning 
how to prepare museum specimens. As I go through 
my routine, I tell them that it is ok to get honest opin¬ 
ions instead of saying that something looks fantastic, 
when it truly looks like it came out of the south end of 
a northbound coyote. 

As time went by, my admiration for Clyde grew 
with my own personal growth in knowledge and where 
I am today as a mammologist. In fact, Clyde’s work 
on bats in New Mexico helped strike a passion in me 
to work on Myotis occultus. I used this passion to help 
resolve a taxonomic question that he had been working 
on with Jim Findley in 1967. Without this inspiration to 
work on M. occultus, I think my MS and PhD degrees 
would have been influenced by another mammal. 


In closing, I must say that in some ways I am 
thankful that I didn’t know who Clyde was when I first 
met him, because I have an unbiased opinion of how 
truly great he was to me and to the field of mammal¬ 
ogy. This became more evident to me as I learned more 
about Clyde through interactions at meetings, social 
gatherings, working together in the field, or from the 
papers he had written. As with many, I think of Clyde 
often and miss him greatly. However, in some ways I 
like to think that he still lives on. I can see this when I 
examine voucher specimens by Clyde that are housed 
at the Museum of Southwestern Biology, but even more 
so when I delve into my fond memories of him and give 
my impersonation of Clyde saying “Valdez, are you 
gonna skin it or make love to it, huh?” as I prepare to 
skin something in front of me. 


Cheri Jones 

Memories of my Dad 


My brother, Craig, and I were born shortly after 
Dad began his work as a graduate student at the Uni¬ 
versity of New Mexico. When we were very small, Dad 
was gone a lot. He was busy with his courses, research, 
and jobs required to support his young family (I seem to 
remember that one of his summer jobs was picking wa¬ 
termelons). We were excited and happy when he came 
home. Mom was the main disciplinarian in those days, 
but Dad mastered the common family strategy known 
as The Look, which I imagine was equally effective 
with errant students later in his career. As he advanced 
after completing his PhD, we moved to New Orleans 
and Mandeville, Louisiana, Bata, Rio Muni (Equatorial 
Guinea), and Vienna, Virginia. We learned to adapt to 
new neighborhoods and new schools fairly quickly. 

Dad kept Walker’s Mammals of the World and 
the Life series of science books at home. However, I 
didn’t immediately gravitate toward biology in public 
school because most of my science classes were less 
than inspirational. One exception was a summer class 
I took in junior high school. That class involved activi¬ 
ties, such as a field trip to a herbarium, and I chose to 
identify and press leaves as my final project. Dad gave 


me my first Peterson field guide, one that addressed the 
wildflowers of the eastern United States. By that time, 
I also was more aware of what fun (and work) was 
involved in biology because of a greater understanding 
of my father’s work. 

Dad and I enjoyed learning and practicing Span¬ 
ish, traveling overseas, and discussing new discoveries 
in zoology. Two special memories involved work as 
field mammalogists. When I started collecting mam¬ 
mals, I discovered that my hands were small enough to 
reach into Sherman traps. Dad’s big hands wouldn’t fit 
inside a live trap, but with those big hands, he prepared 
voucher specimens more quickly and consistently than 
anyone else I’ve ever seen. The second memory I’ll 
recount was when he accompanied me on one of my 
field trips in southern Colorado. After years of hearing 
about his field trips (and going with him on a few), I 
was so excited to have him with me. He really enjoyed 
himself, too; he mentioned that an elk herd we saw was 
the largest he’d ever seen, and he had had little or no 
experience capturing water shrews and jumping mice. 
I really miss Dad, not only as a father, but also because 
I miss sharing so many things with him. 


Encomia and Reflections 


251 


Norm Scott 


Clyde was one of the most important people in my 
life. He took a callow youth who had just lost tenure 
at the University of Connecticut and molded him into 
someone who could contribute meaningfully to the 
scientific community. Early on, he recognized that 
I enjoyed the fieldwork too much and I didn’t follow 
through with the dreary publication process in a timely 
manner; he mentioned it once—and once was enough. 
Clyde also reveled in the fieldwork, and he managed to 
shoehorn it into his heavy administrative and scientific 
schedules whenever he could, and his success at this 
balance provided a model that we all could follow. 

Clyde was a man of bedrock principals, honed 
early in the Sandhills of rural Nebraska. I could always 
depend on him to have my back in the often political 
climate of the Fish and Wildlife Service bureaucracy. 
When one of his team made a mistake that drew the 
attention of the Service, he never even mentioned it to 
us. He took the heat and shielded us, figuring that we 
had already learned a lesson and didn’t need any extra 
pressure. I can’t imagine a better boss, and, when he left 
the Service after a conflict of principles, our tight-knit 
lab unraveled some and was never quite the same again. 

What follows are some of my most valued re¬ 
membrances of Clyde, and what he did with and for 
me. They are in chronological order. 

Barro Colorado Island 

The Howler Monkeys (Alouatta palliata ) at the 
Smithsonian Research Station on Barro Colorado Island 
(BCI) in Gatun Lake, Panama, were one of the most 
studied monkey populations in the world (Carpenter 
1965). In 1973, Richard Thorington, a mammalogist 
at the Smithsonian Museum of Natural History, Wash¬ 
ington D.C., had asked Clyde, then the director of the 
U.S. Fish and Wildlife Service’s National Fish and 
Wildlife Laboratory (NFWL) located in the Museum, 
to recruit a team of biologists to tag Howler Monkeys 
on the island. 

I had been working with a team of graduate 
students at the University of Connecticut, and we 


had developed techniques for capturing and marking 
Howler Monkeys (Scott et al. 1976). We had tagged 
over 100 monkeys in Costa Rica, and Clyde asked me to 
come to BCI to be a part of his team, which included Al 
Gardner and Don Wilson, both NFWL mammalogists. 

Don and I arrived at BCI on the shuttle boat (Fig. 
1), climbed the 350 stairs from Gatun Lake up to the 
station (Fig. 2), and went into the dining room where I 
met Clyde for the first time. He immediately provided 
us all with beer. 


Figure 1. Don Wilson taking boat from Gamboa, Panama, 
to Barro Colorado Island, 1973. 


Figure 2. Smithsonian Tropical Research Institute 
laboratory and dock at Barro Colorado Island on Gatun 
Lake, Panama. 


252 


Clyde Jones Memorial Volume 


We spent the next month tromping over the en¬ 
tire island, catching Howler Monkeys with anesthetic 
darts and tagging them for future studies (Fig. 3). If 
the anesthetic took hold immediately, the monkeys 
fell within 5 or 10 minutes, and we were able to catch 
them in a canvas cloth held up by two people (Fig. 
4). Sometimes, however, the monkey wouldn’t come 
down immediately. They would crawl into the fork of 
the tree, wrap their tail around the branch, and go to 
sleep. Usually, when they began to awaken, they would 
fall out of the fork, but even then they often didn’t lose 
their tail grip, and they would hang head down for up 
to an hour more. Then, when the drug started to finally 
wear off, the monkey would relax its tail and fall. It 


Figure 3. Clyde and Norman taking aim at a Howler 
Monkey. 


Figure 4. Clyde and Norman examining the drugged 
monkey. 


was dangerous for us to stand for more than an hour 
under a 12 kg monkey that could come crashing down 
at any moment, so we piled branches on the ground to 
break the fall and sat down and waited. We weighed 
and measured the drugged monkeys and returned them 
to their home habitat (Fig. 5). 


Figure 5. Clyde ready to return the monkey to its habitat 
and go get another one. 


Old Scar was the dominant male in the troop 
that had its territory around the station. He had been 
named by Carpenter, and was still the lead male when 
we got there. On one of our last days on the island, 
just about lunch time, Clyde and I got a tranquilizer 
dart into Scar’s butt and he hung by his tail. One of 
my most pleasant memories of Clyde was when he 
and I were sitting next to the trail, reminiscing about 
monkeys and waiting for Scar to fall. The crash, when 
it finally came, was a satisfying climax to an unforget¬ 
table month (Fig. 6). 

It wasn’t all monkeys on BCI. To take a break 
on some days, Clyde, with Don and me and Robin An¬ 
drews, a herpetologist, would cruise the forest, looking 
for Noropsfrenatus, a medium-sized lizard that perched 
on tree trunks and searched for invertebrate prey in 
the leaf litter. We developed a theory as to why larger 
lizards perched higher on tree trunks and used larger 
diameter perches (Scott et al. 1976). 


Encomia and Reflections 


253 


Figure 6. Clyde showing old Scar, the lead male in the 
laboratory troop. 


Clyde’s Stryders was a name coined on BCI by 
Thorington (Fig. 7). On Sundays, Clyde’s howler 
monkey team competed with teams of other scientists 
to see who could most rapidly race over the muddy 
forest trails and ascend the 350+ steps that lead from 
Gatun Lake up to the research station. Clyde’s Stryders 
usually took the honors (I remember that Don Wilson 
was the quickest; Fig. 8). 


Figure 7. Clyde and two of his Stryders, Don and Norman. 


Figure 8. Two Stryders, Don and Norman, after the trail/ 
stair mn with a respected opponent from another team. 


University of New Mexico 

After BCI, Clyde hired me to head up a new 
NFWL field station at his alma mater, the University of 
New Mexico. In the first years, I studied (and captured) 
monkeys in Colombia (Scott et al. 1976), Costa Rica 
(Heltne et al. 1976) and Cameroon. 

Clyde’s grant money for monkey research ran 
out in 1976, and I was basically a herpetologist, not a 
primatologist, so, while in Cameroon chasing Black 
Colobus (Colobus satanas) and Gray-cheeked Mang- 
abey (Lophocebus albigena. Fig. 9) monkeys, I returned 
to my roots and extended to Africa previous studies 
that I had done in Costa Rica on the forest leaf litter 
herpetofauna (Fig. 10; Scott 1982). 


254 


Clyde Jones Memorial Volume 


Figure 9. Gray-cheeked Mangabey (Lophocebus 
albigena ), Lombe, Cameroon, 1975. 


Figure 10. Norman with Python regius , Lombe, 
Cameroon. 


Over the next 28 years, the New Mexico NFWL 
team studied vegetation, frogs, toads, turtles, lizards, 
snakes, kangaroo rats, bird faunas, Mexican ducks 
(Anas platyrhynchos diazi) and ants in Texas, Arizona, 
and New Mexico, and in Aguascalientes, Baja Califor¬ 
nia, Sinaloa and Sonora, Mexico. 


In 1987, we resurrected the Stryder team and 
made t-shirts for a 10 km run on the campus of the 
University of New Mexico. Cindy Ramotnik was the 
leading standard-bearer for this event (Fig. 11). 


Figure 11. T-shirt made for the last Stryder mn at the 
University of New Mexico, 1987. 


Isla Tiburon 

Clyde developed a cooperative agreement be¬ 
tween the NFWL lab and Fauna Silvestre, the Mexi¬ 
can equivalent of the U. S. Fish and Wildlife Service. 
Under this agreement, Roy McDiarmid and I gave a 
frog workshop on the shores of Lago Chapala, but the 
main activities were field collecting trips with Mexican 
biologists to Arizona and Baja California. 

On these trips, mammalogists, herpetologists, 
and even sometimes ornithologists, would observe 
and collect specimens of the local faunas, thereby 
greatly extending our knowledge of the abundances 
and distributions of the animals of these biologically 
unexplored regions. Herpetologists were especially 
important on these trips, because the mammalogists 
insisted that we remove all of the rattlesnakes before 
they put out their traplines. 

There are many great memories of these trips, 
but one stands out. In 1976, Clyde, Don Wilson, Mike 
Bogan, and I were on Isla Tiburon in the Sea of Cortes 
(Fig. 12). From our base camp at Caracol (Fig. 13), 


Encomia and Reflections 


255 


we were trapping kangaroo rats (Dipodomys) and other 
small mammals. Above our camp loomed the island’s 
highest point: Cerro Kunkaak (Fig. 14). 


Figure 12. Isla Tiburon, Sonora, Mexico. Solid lines are 
roads, dashed line is the hike that Clyde and I took. 


Figure 13. Clyde in camp at Caracol, Isla Tiburon, Mexico, 
1976. 


Figure 14. Cerro Kunkaak at dawn, Isla Tiburon, Mexico. 


After an overnight trapping session at a spring 
in Arroyo Chalate on the island’s east coast, Mike and 
Don left in the truck, and Clyde and I started to hike 
back to our camp at Caracol. We knew what direction 
the camp was, so we headed across the island on foot. 
However, we made the mistake of heading straight for 
Caracol instead of going around the mountain range 
in between. The terrain was rough and steep, and we 
followed canyons and ridge lines, going ever higher. 
We ended up spending the afternoon napping in a cave 
near Cerro Kunkaak. In the cave, Clyde found the skull 
of a ring-tailed cat (Bassariscus astutus ) which is now 
in the National Museum (USNM 514028). This, and 
another skull (USNM 514027) from the same trip, may 
be the only records for the species from the island. We 
had only brought a liter of water with us, and, by this 
time, we were getting pretty dry. 

After our nap, thirsty and blocked by the vertical, 
colorful red and green lichen-covered walls of Cerro 
Kunkaak (Fig. 15), we headed down the nearest canyon. 
Lo and behold, we were saved when we came upon a 
large tinaja, a 3m x 18m pool of water in the eroded 
igneous rocks of the canyon floor (Fig. 16). 


Figure 15. Summit of Cerro Kunkaak, Isla Tiburon, 
Mexico. 


Figure 16. Life-saving tinaja (waterhole) with fig tree, 
Isla Tiburon, Mexico. 


256 


Clyde Jones Memorial Volume 


By this time it was getting late, and we hiked 
down the canyon to the bajada at the base of the 
mountain at night, and there we slept until dawn. In 
the morning, we started walking around the base of the 
mountain to the road to Caracol, where we met a search 
party that was looking for us (Fig. 17). 


Figure 17. Central IslaTiburon, Sonora, Mexico. Solid 
lines are roads, dashed line is the hike that Clyde and 
I took. Map courtesy of Google, Terrametrics, Digital 
Globe and INEGI; data from SIU, NOAA, U.S Navy, 
NGA andGEBCO. 


Colorado River 

I think that Clyde’s favorite professional activity 
was the raft trips that he organized on the Colorado Riv¬ 
er through the Grand Canyon. In 1969, he left Tulane 
University, and in 1970, he went to Washington D.C. 
to be the Chief of the U.S. Department of the Interior’s 
Bird and Mammal Laboratories located in the National 
Museum. Clyde developed a project with the National 
Park Service to survey and collect the small vertebrates 
present along and in the Colorado River through the 
Grand Canyon. From 1970 through 1981, he went on 
at least 16 river trips, many of them covering the entire 
290 miles from Lee’s Ferry to Lake Mead (Fig. 18). 
The trips included up to 17 people, mostly scientists, 
but also some “significant others” that went along for 
support. They took habitat photographs in canyons 
surrounded by towering cliffs (Fig. 19), and they ran 
river rapids (Fig. 20). During afternoons and nights 
ashore, they made collections of plants, fish, small 
mammals, reptiles and amphibians. The 14 trips from 
1970 to 1976 resulted in a report to the Park Service 
(Suttkus et al. 1976). 


Figure 18. Start of Colorado River trip, Lee’s Ferry, 
Arizona, 1980. 


Figure 19. Colorado River, Fern Glen Canyon. 


Figure 20. Colorado River, Hermit Rapids. 


For fish studies, he recruited two ichthyologists, 
former colleagues from Tulane University, Royal 
Suttkus (Sut) and his student, Glenn Clemmer (then 
at Mississippi State University). They were the core 


Encomia and Reflections 


257 


team that was surveying (Fig. 21) for any remnant 
populations of six native fish species that had been 
largely eliminated from the main Colorado River by 
coldwater releases from the bottom of the lake behind 
Glen Canyon Dam. Small populations of the Federally 
Endangered Humpback Chub ( Gila cypha. Fig. 22), 
the Flannelmouth Sucker ( Catostomus latipinnis ), the 
Bluehead Sucker ( Catostomus discobolus ), and the 
Speckled Dace ( Rhinichthys osculus) persisted in the 
wanner waters at the mouths of the larger tributaries to 
the river; however, two other native species, the endan¬ 
gered Colorado Squawfish ( Ptychocheilus lucius ) and 
the endangered Razorback Sucker ( Xyrauchen texanus ) 
were not found and are believed to have been extirpated 
from the river in the Grand Canyon. The team made 
several recommendations for the management of the 
critically endangered Humpback Chub (Suttkus et al. 
1976; Valdez and Clemmer 1982). 


Figure 21. Clyde, Glenn, and Sut seining at the mouth of 
the Little Colorado River. 


Figure 22. Humpback Chub ( Gila cypha ) in net, Little 
Colorado River. 


Between the fish sampling sites, Clyde and vari¬ 
ous mammalogists and herpetologists set out traplines 
and hunted for specimens. By the time the trips ended 
in 1981, the project had recorded 19 species of fish, 
four amphibians, 16 reptiles, and 28 mammals from the 
river and the canyon floor (Suttkus et al. 1976, 1978). 

Joan and I participated in the last Colorado River 
trip. The boat ran out of gas when we reached the upper 
reaches of Lake Mead, and we spent the eve of our 25 th 
wedding anniversary floating at the whim of the gentle 
breezes. By this time on the trip, alcohol was getting 
scarce, but Pat Mehlhop had brought a bottle of Lejon 
Cold Duck champagne just for the occasion, and we 
had saved a bottle of Kahlua. I spent the evening in the 
warm waters of Lake Mead with a bottle, going around 
the raft filling everyone’s cups as I went, all except for 
Clyde, who was still drinking beer. It was a unique 
celebration to say the least. 

For Clyde, those trips were more than collecting 
expeditions—they were a spiritual experience, facili¬ 
tated by the absence of ringing phones and bureaucratic 
meetings, and surrounded by friends, all working to¬ 
wards the same goal, all the time overshadowed by 
the towering cliffs and hanging gardens of the canyon 
(Fig. 23). One of my fondest memories of Clyde is of 
him standing in the bow of the raft, gazing up at the 
looming walls, and hearing him say that these trips were 
the closest that he had ever felt to having a religious 
experience. 


Postlude. —Clyde was a wonderful boss, col¬ 
league and friend, and I miss him greatly. 


Figure 23. Colorado River; Clyde and colleagues at the 
mouth of Havasu Creek. 


258 


Clyde Jones Memorial Volume 


Literature Cited 


Carpenter, C. R. 1965. The howlers of Barro Colorado Island. 
Pp. 250-291 in Primate Behavior (I. DeVore, ed.). Holt, 
Rinehart and Winston, New York. 

Heltne, P. G., D. C. Turner, andN. J. Scott, Jr. 1976. Compari¬ 
son of census data on Alouatta villosa (= pallicita) from 
Costa Rica and Panama. Pp. 10-19 in Neotropical Pri¬ 
mates: Field Studies and Conservation (R. J. Thorington 
and P. G. Heltne, eds). National Academy of Sciences, 
Washington, DC. 

Scott, N. J., Jr., D. E. Wilson, C. Jones, and R. M. Andrews. 
1976. The choice of perch dimensions by Anolis lizards. 
Journal of Herpetology 10:75-84. 

Scott, N. J., Jr., A. F. Scott, and L. A. Malmgren. 1976. Captur¬ 
ing and marking howler monkeys (Alouatta pallicita ) for 
field behavioral studies. Primates 17:527533. 

Scott, N. J., Jr., T. T. Struhsaker, K. Glander, and H. Chirivi. 
1976. Primates and their habitats in northern Colom¬ 
bia with recommendations for future management and 
research. Pp. 3050 in First Inter-American Conference 
on Conservation and Utilization of American Nonhuman 
Primates in Biomedical Research. Pan American Health 
Organization Scientific Publication 317. 

Scott, N. J., Jr. 1982. The herpetofauna of forest litter plots from 
Cameroon. Pp. 145-150 in Herpetological Communities: 


A Symposium of the Society for the Study of Amphibians 
and Reptiles and the Herpetologists’ League, August 1977 
(N. J. Scott, Jr., ed.). Fish and Wildlife Service, Wildlife 
Research Report 13. 

Suttkus, R. D., and G. H. Clemmer. 1977. The humpback chub, 
Gila cypha, in the Grand Canyon area of the Colorado 
River. Occasional Papers, Tulane University Museum of 
Natural History 1:1-30. 

Suttkus, R. D, G. H. Clemmer, C. Jones, and C. R. Shoop. 
1976. Survey of fishes, mammals and herpetofauna of 
the Colorado River in Grand Canyon. Grand Canyon 
National Park, Colorado River Research Series, Contri¬ 
bution 34:1-48. 

Suttkus, R. D, G. H. Clemmer, and C. Jones. 1978. Mammals 
of the riparian region of the Colorado River in the Grand 
Canyon area of Arizona. Occasional Papers, Tulane 
University Museum of Natural History 2:1-32. 

Valdez, R. A, and G. H. Clemmer. 1982. Life history and pros¬ 
pects for recovery of the humpback and bonytail chub. 
Proceedings of a Symposium on Fishes of the Upper 
Colorado River System: Present and Future. American 
Fisheries Society, Bethesda, Maryland. 


Photographs 


Photo 1. Clyde with pygmoid forest dwellers, Rio Muni, West Africa. Photo submitted by 
David Marshall. 


Photo 2. Footbridge. West African rain forest, Rio Muni, 1967. Photo submitted by Mary 
Ann Jones. 


259 


260 


Clyde Jones Memorial Volume 


Photo 3. Habitat shot, Rio Muni, 1968. Photo submitted by Mary Ann Jones. 


Photo 4. Making arrangements for lodging and a tracker. Evinayong, Rio Muni, 1968. Photo 
submitted by Mary Ann Jones. 


Photographs 


261 


Photo 5. Receiving an Antarctic medal, 1971. Photo 
submitted by Mary Ann Jones. 


Photo 6. Clyde (back row, right), Royal Suttkus (front row, fourth from left) and the rest of the 
‘River Rats’, Colorado River, ca. 1970s. Photo submitted by Mary Ann Jones. 


262 


Clyde Jones Memorial Volume 


Photo 7. “All in a Day’s Work.” Depicted (left to right) are: Rick Manning, Clyde Jones, J. 
Knox Jones, Jr., and Larry Choate. Mammals were taken in Andrews and Winkler counties, 
Texas (ca. Spring, 1989). Photo taken at the NSRL, mammal prep room. Photo submitted 
by Rick Manning. 


Photo 8. “Inspecting the rats.” J Knox Jones, Jr. and Clyde Jones with mammals taken at Harte 
Ranch, Brewster County, Texas (ca. October 1991) by Rick Manning and Jim Goetze. Photo 
taken at the NSRL, mammal prep room. Photo submitted by Rick Manning. 


Photographs 


263 


Photo9. Seining for fish, Limpia Creek, Fort Davis, 1998. Photo submitted by Mary Ann Jones. 


Photo 10. Collecting fish from a seine, Limpia Creek, Fort Davis, 1998. Photo submitted by 
Mary Ann Jones. 


264 


Clyde Jones Memorial Volume 


Photo 11. Clyde at Big Bend Ranch State Park, 1998. Photo submitted by Mary Ann Jones. 


Photo 12. Ron Van Den Bussche, Robert Bradley, Meredith Hamilton, and Clyde preparing 
beer-batter pancakes in the Davis Mountains, 1998. Photo submitted by Robert D. Bradley. 


Photographs 


265 


Photo 13. Clyde preparing a specimen in the Davis 
Mountains, 1999. Photo submitted by Robert D. Bradley. 


Photo 14. Sowell Expedition, Ecuador, 2004. Photo submitted by Lisa Bradley. 


266 


Clyde Jones Memorial Volume 


Photo 15. Clyde measuring a mouse. Photo submitted by David 
Riskind. 


Photo 16. David Riskind and Clyde, San Antonio Cabin, Chinati Mountains State Natural Area, 
2005. Photo submitted by David Riskind. 


Photographs 


267 


Photo 17. Mark Lockwood and Clyde at a field camp in Teneros Creek, Big Bend Ranch State 
Park, 2002. Photo submitted by David Riskind. 


Photo 18. Clyde landing on top of Sierra Parda, Chinati Mountains State Natural Area, 2005. 
Photo submitted by Frank Yancey. 


268 


Clyde Jones Memorial Volume 


Photo 19. Mike Bogan, Cindy Ramotnik, Clyde, and Frank Yancey on a collecting trip to the 
Chinati Mountains, 2005. Photo submitted by Frank Yancey. 


Photo 20. Clyde, Mike Bogan, A1 Gardner, and Robert Fisher. 83rd Annual Meeting of the 
American Society of Mammalogists, hosted by Texas Tech University, 2003. Photo submitted 
by Mary Ann Jones. 


Photographs 


269 


Photo 21. A moment of reflection after preparing several bat specimens - each one a “thing of 
beauty and joy forever”. San Antonio Cabin, Chinati Mountains, 2007. Photo submitted by 
Mary Ann Jones. 


Photo 22. Clyde with colleagues, friends, and former students. Standing, left to right: Steve 
Kasper, Joel Brant, Rick Manning, and Larry Choate. Seated, left to right: Jim Goetze, Frank 
Yancey, Clyde Jones, and Fred Stangl. Texas Society of Mammalogists, Junction, Texas, 2013. 
Photo submitted by Lisa Bradley. 


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Clyde Jones Memorial Volume 


Photo 23. Sherman trap damaged by hail. Big Bend Ranch State Park, June 2013. Photo 
submitted by Mary Ann Jones. 


Photo 24. Clyde with former students (and the editors of this volume) Rick Manning, Frank 
Yancey, and Jim Goetze. Big Bend Ranch State Park, June 2013. Photo submitted by Mary 
Ann Jones. 


Photographs 


271 


Photo 25. Clyde with part of the Big Bend Ranch State Park field crew, June 2013. Photo 
submitted by Mary Ann Jones. 


Photo 26. Clyde, Mary Ann, Erica, and Lou at the Texas Tech Frazier Alumni Pavilion, 2013. 
Photo submitted by Lou Densmore. 


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Clyde Jones Memorial Volume 


Photo 27. Clyde at TSM meeting, 2014. Photo submitted by 
Lisa Bradley. 


Photo 28. Rick Manning, Clyde, and Frank Yancey, Junction, Texas (post-TSM meeting), 
February 2015. Photo submitted by Frank Yancey. 


Photographs 


273 


In Memory 


Addendum 


Editors ’ note: Due to various factors, this encomium was submitted too late to be included in the published 
memorial volume. However, in consideration of the relationship of the authors to Clyde, we agreed to make this 
encomium available via electronic format as a supplement to the printed volume. 


Robert J. Baker, Carleton J. Phillips, and Hugh H. Genoways 


This is our collective attempt to provide an en¬ 
comium (an honest song of praise) for Clyde Jones. 
After some urging from the editors of this volume, we 
conspired to share some of our memories of Clyde. 
We preface this effort by noting two things: 1) this 
is our second attempt at writing an encomium for a 
departed colleague; and 2) that we three each became 
friends individually with Clyde when we were young 
(child-like) mammalogists attempting to establish 
ourselves as scholars and also as members of a team 
of people who would be field biologists following in 
the traditions of such mammalogical luminaries as C. 
Hart Merriam, Hartley H. T. Jackson, Joseph Grinnell, 
E.A. Goldman, GerritS. Miller, Jr., EmmetT. Hooper, 
James S. Findley, Rollin H. Baker, J. Knox Jones, Jr., 
Sydney Anderson, and E. Raymond Hall. 

Encomia can be tough, because although they 
are odes of praise, they are expected to be honest ap¬ 
praisals. If that kind of complexity gives you pause, 
then just think of Marc Antony’s encomium for Caesar! 
Sometimes taking an honest approach ends up being a 
“warts and all” outcome to telling about the person in 
question. But sometimes it does not take such a path. 
Inevitably, readers (who have their own opinions and 
always think correctly that they would have written the 
encomium differently) are most likely to come away 
with a bad taste in their mouth. But the thing of it is that 
encomia, if they are honest, often tell the reader more 
about the author(s) and their knowledge of the principal 
than they tell about the principal himself. It is impos¬ 
sible to write any encomium without self-revelation. 
Our first attempt at this was with a volume of articles 
in honor of J. Knox Jones, Jr. (Genoways, H. H., and 
R. J. Baker [eds.]. 1996. Contributions in mammal¬ 
ogy: a memorial volume honoring Dr. J. Knox Jones, 
Jr. Museum of Texas Tech University, Lubbock, il + 
315 pp.). We asked all authors to write an encomium 
for Knox, and the results were lor 2 mm from disaster. 
What we did was unique, scary, and not so nice, but in 
the name of honesty Knox deserved it. Clyde, on the 
other hand, does not deserve it but that alone is a huge 


complement. It is fitting that Clyde and Knox were 
close friends (“buds” is the term they used). Knox took 
advantage of Clyde at every opportunity, just as he did 
with everyone else in his life. 

We each had our own individual relationship 
with Clyde, which is something we discovered when 
we tried to write this encomium collectively rather 
than as individuals. Robert Baker recalls his first meet¬ 
ing with Clyde Jones was at the American Society of 
Mammalogists (ASM) meetings in Long Beach in 1966 
where Clyde was one of the first platform speakers at 
that meeting. The science of mammalogy at that time 
was mostly about fieldwork, but Baker remembers that 
Clyde talked about his ecological research on bats in 
New Mexico. Clyde’s later work on the Rio Muni pri¬ 
mate fauna was incredibly powerful. Baker recollects 
fantasizing about getting to go to the field in this Afri¬ 
can location and getting to collect primates and bats. 
When Clyde told stories about these field endeavors, it 
evoked jealousy in Baker because his own background 
had been fieldwork in Arkansas, Arizona, and Mexico. 
Over the next several years Clyde was always present 
at the mammal meetings and he usually had a bar in his 
room, primarily with bourbon as Baker recalls. Late in 
his life, Clyde replaced the bourbon with cheap (awful) 
Texas canned beer that he diluted with ice. 

Baker recollects conversations with Hugh Ge¬ 
noways early in their involvement in the ASM about 
Clyde’s tremendous mental organization and seem¬ 
ingly global knowledge about mammalogy. Baker 
also recalls expecting Clyde ultimately to provide 
leadership to the ASM. As Baker remembers, Don 
Wilson shared his belief and also was very prominent 
in those early discussions. The fact that both Clyde 
and Don Wilson were associated with the U.S. Fish 
and Wildlife Service housed at the National Museum 
of Natural History gave these two young mammalo¬ 
gists instant credibility among their contemporaries. 
Robert’s premonition that Clyde would assume lead¬ 
ership roles in the ASM was prophetic because Clyde 


2 


Clyde Jones Memorial Volume 


served on 10 Society committees: Index (1972-1974); 
Editorial (1972-1976, 1984-1990); Conservation of 
Land Mammals (1973-1974); Information Retrieval 
(1972-1979); Systematics Collections (1972-1986); 
C. Hart Merriam Award Committee (1974); Legisla¬ 
tion and Regulations (1975-1979, 1990-1992); Hart¬ 
ley H. T. Jackson Award (1977-1978); Development 
(1988-1990); and Nomenclature (1995-1998). Clyde 
also served the ASM in a number of other capacities, 
as well, including as a member of the Board of Direc¬ 
tors (1971-1979, 1984-1990), Editor for Reviews 
(1972-1976), and Managing Editor for the Journal of 
Mammalogy (1984-1990). His work on behalf of the 
ASM was recognized with the Hartley H. T. Jackson 
Award “for long and outstanding service to the ASM” in 
1997 and Honorary Membership “conferred in recogni¬ 
tion of distinguished service to mammalogy” in 2003. 
In other organizations, Clyde served as President of the 
Biological Society of Washington in 1975 and President 
of the Texas Society of Mammalogists in 1987. 

Thinking back, Baker was particularly excited 
when Clyde became Director of the Museum of Texas 
Tech University in 1982 and later was active in the 
Department of Biological Sciences. It gave Baker his 
first opportunity to interact with Clyde on a daily basis 
and to find his first impressions were more than correct. 
It was Baker’s pleasure and honor to initiate the effort 
in 1999, along with Carl Phillips who was Department 
Chair at the time, that resulted in successfully getting 
Clyde appointed as a Paul Whitfield Horn Professor 
(named in honor the first president of TTU), which is 
the highest honor bestowed on a Texas Tech faculty 
member. This recognition is given to a faculty member 
who has attained national and international prominence 
in the area of his research. 

In contrast to Baker, Phillips has no early—gradu¬ 
ate school—personal memories of Clyde. However, it 
seems likely that the two of them first became acquaint¬ 
ed at the ASM meeting held in Pacific Grove, Califor¬ 
nia, in 1973. At that meeting, Phillips remembers that 
the Journal of Assinology [a spoof journal] made one 
of its last unauthorized appearances, with a drawing of 
a naked Clyde as a centerfold—a male parody of what 
Playboy magazine made popular. That particular issue 
of Assinology , which was created largely by James 
Dale Smith, marked the perigee (or apogee, depending 
on one’s opinion of such things) of its transformation 


from cute and witty (at least to insiders) to gross and 
embarrassing to all. 

Ironically, thanks to their meeting at the time of 
Clyde’s unwanted appearance in the Journal of Assinol¬ 
ogy, Clyde and Carl knew each other professionally for 
nearly 40 years before they unexpectedly found them¬ 
selves working together. When Phillips was recruited to 
Texas Tech University as Chair of Biological Sciences 
in 1998, Clyde agreed to serve as one of his Associate 
Chairs. Choosing Clyde was a classic “no-brainer.” 
Clyde had served as a faculty member and as Director 
of the Museum of Texas Tech University. He had an 
encyclopedic grasp of the Department (a very large 
academic unit with 40 faculty with diverse interests), 
the university, and assorted personalities important to 
the daily life of the Department, including the Provost 
(John Burns) and Vice President for Research (David 
Schmidly), both of whom were mammalogists. From 
his Fish and Wildlife days, Clyde had acquired political 
and corporate experience far beyond that of a typical 
university professor and that fact, along with his time 
in service, (he knew where the bodies were buried) 
made him valuable to Phillips. Clyde was skilled 
as an Associate Chair. He was particularly good at 
semi-secret diplomatic missions on behalf of Phillips. 
This especially was the case with the Provost, who 
had been a faculty member and Chair of Biological 
Sciences—the worst-case scenario for any incoming 
new Chair. Whenever the Provost’s meddling in de¬ 
partmental business became too much to bear, Phillips 
would send Clyde on a mission to visit the Provost and 
cool off the meddling behavior. Beyond this sort of 
special work, Clyde’s loyalty to the Department was 
well known and widely appreciated. In 2001, Phillips 
and Baker obtained funding for fieldwork in Ecuador. 
Clyde visited with the two of them and requested an 
invitation to join them and help with the collecting 
project. By that time he had done fieldwork on every 
continent except South America, so naturally he was 
anxious to complete his list of continents. This was 
the first time that Phillips had conducted fieldwork 
with Clyde Jones: the surprise—or not so much of a 
surprise—was that Clyde was strictly old school. So, 
while Phillips and Baker prepared specimens for future 
genomic studies, transmission electron microscopy, and 
in situ chromosomal hybridizations, Clyde strictly did 
museum preparations following protocols popular in 
1895, and even earlier. As it turned out, there was no 


Addendum—Encomia and Reflections 


3 


way to dissuade him from what seemed to Phillips and 
Baker as a wasteful approach. The waste was due to 
loss of potential data—once a mammal is euthanized 
retrieval of tissues has to be immediate. On the posi¬ 
tive side, Clyde’s personality was such that he worked 
very well with the American and Ecuadoran students 
participating in the field project. Teaching traditional 
field mammalogy was Clyde’s big contribution to the 
program. And the same was true back on campus with 
Clyde’s graduate students. 

One evening, Carl Phillips and Clyde Jones 
visited over scotch (Phillips) and awful beer with ice 
cubes (Jones), while discussing the history of American 
mammalogy, and in particular its most likely future 
directions, they hatched an idea that began with a ques¬ 
tion. Why not assemble a group of prominent scientists 
with similar backgrounds but diverse interests and ask 
them to talk and write about their professional lives 
and future of the discipline? Such a project would be 
unique. The next goal was to fund and host a confer¬ 
ence followed by a publication of autobiographies that 
would be valuable to future students and science histo¬ 
rians alike. The only criterion for participation was that 
the mammalogist had to have had at least 30 years of 
experience in his or her field. It quickly became clear 
that the most logical candidates shared academic ties 
that reflected the origin and history of North American 
mammalogy. Phillips and Jones edited the volume, 
entitled Going Afield, which can be described as suc¬ 
cessful (and now out of print). 

Hugh Genoways does not recall when he first met 
Clyde in person, but he had already heard about Clyde 
and their shared Nebraska roots. The first meeting 
would have been in the Mammal Range at the National 
Museum of Natural History when Hugh was visiting 
to study his beloved Liomys or at an ASM meeting 
as they became more involved in the activities of the 
Society. Whenever that meeting occurred there was 
an immediate connection based on a mutual love of 
mammals, fieldwork, and Nebraska. Over the years, 
they met on many different occasions, but really only 
worked together in editing a book on museological 
practices for mammal collections. As time moved 
along Hugh became more and more aware of Clyde’s 
biography and was always struck by how much they 
had in common. The full extent of this common bi¬ 
ography only became obvious with the publication of 


Clyde’s autobiographical piece “You Have to Catch 
Them First” in Going Afield (Museum of Texas Tech 
University, 2005). For example, Clyde was born in 
Scottsbluff, Nebraska, on 3 March 1935, whereas Hugh 
was born in the same hospital (there was only one) 
on 24 December 1940. Although Clyde’s family left 
western Nebraska, they were engaged in farming and 
ranching near Burwell, NE, (pop. 1,400 in 1940) on 
the eastern edge of the Nebraska Sandhills, Hugh and 
his family remained in America’s Valley of the Nile, 
the North Platte River Valley, living around and finally 
in the town of Bayard (pop. 2,000 in 1940), where his 
father was a farmer and then worked for the Great West¬ 
ern Sugar Company (turning sugar beets into sugar). 
Their mothers were both schoolteachers. They both 
began, and survived, educations in one-room country 
schools. Clyde’s graduating high school class was 32 
(unfortunately, Clyde was not there in the fall of 2016 
when his Burwell Fonghoms won their first Nebraska 
State Championship in class D1 football) and Hugh’s 
was 44. Upon graduation, Clyde enrolled in Hastings 
College (Hastings, NE) in 1953, graduating in 1957. 
Hugh, upon high school graduation, entered Hastings 
College in 1959, graduating in 1963. They took many 
of the same courses from the same professors. Clyde 
played football with considerable success (his 1954 
team was elected to the Hastings College Athletic Hall 
of Fame in 1989), but gave it up for academics in his 
senior year, whereas Hugh lasted only one year in the 
football program with the same coach before discover¬ 
ing Comparative Anatomy and the challenges of schol¬ 
arly pursuits. Clyde made a life-long friend at Hastings 
in Eugene Fleharty, another Nebraska boy who went 
on to dedicate much of his life to studying mammals 
and the Great Plains. Hugh found a fellow traveler 
in Charles Fowler at Hastings, another Nebraska boy, 
whose professional pursuits ultimately involved studies 
of northern fur seals for NOAA. Clyde went off to the 
University of New Mexico for his graduate education, 
whereas Hugh went to the University of Kansas to 
work with yet another Nebraska native, J. Knox Jones, 
Jr. Finally, although their times did not overlap, they 
both served on the faculty at Texas Tech University 
and worked for the Museum of Texas Tech University. 

Nebraska is a large state in area, but less than 
1.5 million people were scattered over these distances. 
What were (or are) the shared experiences and back¬ 
grounds that led so many Nebraskans to the study of 


4 


Clyde Jones Memorial Volume 


modern mammals? Clyde’s autobiography certainly 
makes some of these factors quite clear—the love of 
empty places (Hugh calls it the high lonesome); love of 
the land and landscapes; working with, and knowledge 
of, animals; a general understanding of the environment 
and how it works; ability to work alone; love of being 
out of doors; and self-motivation. Many probably were 
motivated also by the thought of finding a way to get 
out of Nebraska. Nebraska has also produced its share 
of ornithologists and herpetologists and certainly more 
than its share of vertebrate paleontologists, but here is 
Hugh’s admittedly incomplete list (other than those 
listed above) of native Nebraskans who have shared 
Clyde’s and Hugh’s experiences over the generations: 
Lawrence Bruner, Merritt Cary, Melbourne A. Carriker, 
Myron Swenk, Robert Packard, William F. Andelt, Rob¬ 
ert M. Timm, John Comely (another Hastings College 
graduate), Richard W. Manning, Keith Geluso, Jeffrey 
Huebschman, Teresa Zimmerman Frink, Justin Hoff¬ 
man, and Zachary Roehrs. If you don’t know some of 
these people, you should. 

Hugh was always impressed with what Clyde 
was able to accomplish in 20 months in Rio Muni 
[now Equatorial Guinea] in 1966-1968. This work 
was carried out under field conditions that could only 
be described as “primitive.” No one was there to watch 
him or to be certain that he conducted his research. The 
information that he gathered during this work resulted 
in 15 of his first 50 publications, covering his target 
primates as well as information about rodents and 
bats. His classic work “Comparative ecology of Go¬ 
rilla gorilla (Savage and Wyman) and Pan troglodytes 
(Blumenback) in Rio Muni, West Africa,” published 
with his co-investigator Sabater Pi in Bibliotheca 
Primatologica (13:iv + 96 pp., 1971) stands as the 
seminal work on lowland gorillas. This comparative 
study of these two great apes remains unmatched in the 
primatological literature. 

One of our co-author colleagues seems to be 
somewhat critical of Clyde; stating that he “strictly did 


museum preparations following protocols popular in 
1895 . . . there was no way to dissuade him from what 
seemed to Phillips and Baker as a wasteful approach.” 
Hugh is certain that much the same things are said 
about him, when he is not listening, so that a defense 
of these practices seems appropriate at the end of this 
“honest song of praise.” Thank God! Someone has 
been around since 1895, and before, who was willing 
to take the time and effort to do museum preparation of 
mammals. Where would our science of mammalogy 
be without those efforts? How would we anchor our 
knowledge about species without museum vouchers? 
The generation of mammalogists who were trained 
in more traditional practices of mammalogy, but later 
have taken up genomic studies, transmission electron 
microscopy, and in situ chromosomal hybridizations 
during their careers have understood the need for 
voucher specimens for their research. Unfortunately, to 
the detriment of our science, the present generations of 
researchers pursuing these and related studies have not 
been trained in these traditions. Mammals are known 
to these young researchers as pieces of frozen tissue, 
or cell lines, or genetic sequences from GenBank®. 
Little or no thought is given to these resources or the 
time and effort that it has taken to gather, preserve, 
and document them, but rather these are treated like 
a birthright. Seldom are the sources or the scientists 
involved in building the foundation of these resources 
even acknowledged. Clyde and Hugh would give 
these young scientists the same advice that Jim Findley 
gave Clyde: “You have to catch them first.” You must 
become familiar with the whole mammal functioning 
in its environment. You must become a contributor to 
replenish the resource that you are using, be it frozen 
tissue or cell lines or any other stored resource. You 
must become aware of the value of voucher specimens 
to help place the results of your studies into the context 
of the science of mammalogy.