Resources U.S. Department of the Interior U.S. Geological Survey Southwest Biological Science Center Open-File Report 2005-1141 April 2005 In Cooperation with the University of Arizona School of Natural and Arizona Game and Fish Department Water Requirements of Desert Ungulates Water Literature Review and Annotated Bibliography: g r a p n s& t o T c u k r n e r v i e w a n C ain III, Kra usm an, Rose Literature R e d A nnotated Biblio O pen-File Report 2005-1141 hy: W ater Require m ents of D esert U ngulates In cooperation with the University of Arizona School of Natural Resources and Arizona Game and Fish Department Literature Review and Annotated Bibliography: Water Requirements of Desert Ungulates By James W. Cain III, Paul R. Krausman, Steven S. Rosenstock, and Jack C. Turner Open-File Report 2005-1141 April 2005 USGS Southwest Biological Science Center Sonoran Desert Research Station University of Arizona U.S. Department of the Interior School of Natural Resources 125 Biological Sciences East U.S. Geological Survey Tucson, Arizona 85721 U.S. Department of the Interior Gale A. Norton, Secretary U.S. Geological Survey Charles G. Groat, Director U.S. Geological Survey, Reston, Virginia: 2005 Note: This document contains information of a preliminary nature and was prepared primarily for internal use in the U.S. Geological Survey. This information is NOT intended for use in open literature prior to publication by the investigators named unless permission is obtained in writing from the investigators named and from the Station Leader. Suggested Citation Cain III, J.W., P.R. Krausman, S.S. Rosenstock, J.C. Turner. 2005. Literature Review and Annotated Bibliography: Water Requirements of Desert Ungulates. USGS Open-File Report 2005- 1141. U.S. Geological Survey, Southwest Biological Science Center, Sonoran Desert Research Station, University of Arizona, Tucson, AZ. Cover photo: James W. Cain III, Sierra Pinta Mountains, Cabeza Prieta National Wildlife Refuge Authors James W. Cain III Steven S. Rosenstock University of Arizona Arizona Game and Fish Department School of Natural Resources Research Branch 104 Biological Sciences East 2221 W Greenway Road Tucson, Arizona 85721 Phoenix, Arizona 85023 Paul R. Krausman Jack C. Turner University of Arizona Sam Houston State University School of Natural Resources P.O. Box 2116 325 Biological Sciences East Huntsville, Texas 77341 Tucson, Arizona 85721 U.S. Geological Survey Station Personnel Charles Van Riper, III, Station Leader Steve P. Gloss, Ecologist William L. Halvorson, Research Ecologist Cecil R. Schwalbe, Ecologist Michael R. Kunzmann, Ecologist Program and Expertise Areas of USGS and UA Personnel Administration & Outreach Fire Management Cori Dolan Cori Dolan Wendy Parrish James Feldmann Charles Van Riper, III Bill Halvorson Avian Ecology Invasive Species Research Claire Crow Patricia Guertin Glenn Johnson Jim Malusa Mike Kunzmann Phil Rosen Chris O’Brien Cecil Schwalbe Brian Powell Brent Sigafus Charles van Riper, III Dennis Suhre Data Management Inventory & Monitoring Mike Kunzmann Patricia Guertin Brent Sigafus Bill Halvorson Brian Powell Ecology of Amphibians & Reptiles Cecilia Schmidt Kevin Baker Cristina Jones Vegetation Mapping & Ecology Dave Prival Patricia Guertin Phil Rosen Bill Halvorson Cecil Schwalbe Jim Malusa Brent Sigafus Eric Stitt Dennis Suhre USGS Southwest Biological Science Center http://sbsc.wr.usgs.gov Table of Contents Executive Summary......................................................................................................................................1 Introduction and Definitions .........................................................................................................................1 Behavioral Adaptations ............................................................................................................................2 Morphological Adaptations ......................................................................................................................3 Physiological Adaptations........................................................................................................................4 Methods for Studying Water Physiology......................................................................................................9 Water Balance and Water Flux.................................................................................................................9 Blood Constituents .................................................................................................................................10 Water Loss..............................................................................................................................................10 Literature Cited ...........................................................................................................................................11 Annotated Biblography...............................................................................................................................19 Ungulate Mammals ................................................................................................................................19 Marsupial Mammals...............................................................................................................................86 Small Eutherian Mammals .....................................................................................................................92 Techniques and Methodology ..............................................................................................................108 General Desert Ecophysiology .............................................................................................................110 Acknowledgements This study was funded and supported by the Arizona Game and Fish Department, Research Branch and the School of Natural Resources, The University of Arizona. iv Executive Summary in time. Water flux or water turnover is the amount of water (ml/day) an animal processes Ungulates adapted to desert areas are able to through its body each day (Nagy and Peterson survive extreme temperatures and limited water 1988). availability. This ability is largely due to behavioral, morphological, and physiological We review literature relating to the effects of adaptations that allow these animals to avoid or dehydration on wild and domestic ungulates and tolerate extreme environmental conditions. The also discuss the behavioral, morphological, and physiological adaptations possessed by ungulates physiological mechanisms used by ungulates to for thermoregulation and maintenance of water maintain water and temperature balance in an arid balance have been the subject of numerous studies environment: involving a wide range of species. In this report we review the behavioral, morphological, and Behavioral adaptations reviewed include physiological mechanisms used by ungulates and timing of activities, use of microhabitats, other desert mammals to maintain water and body orientation, migration, diet selection, temperature balance in arid environments. We and social behaviors. also review some of the more commonly used Morphological adaptations include body size methods for studying the physiological and shape, pelage and skin characteristics, and mechanisms involved in water balance and patterns of fat deposition. thermoregulation, and the influence of Physiological adaptations include adaptive dehydration on these mechanisms. heterothermy and selective brain cooling, as well as adjustments to metabolic rate, renal and digestive water loss, and pulmonary and Introduction and Definitions cutaneous evaporative water loss. For this literature review and annotated Finally, we review some commonly used methods bibliography our objectives were (1) to compile for the study of the physiological mechanisms available scientific literature on thermoregulation involved in water balance and thermoregulation, and water balance in desert mammals, particularly and the influence of dehydration on these studies on ungulates and (2) to review the mechanisms. We have included some studies methodologies available to study the conducted on the physiology of thermoregulation physiological mechanisms involved in and water balance on small eutherian and thermoregulation and water balance. We were marsupial mammals. specifically interested in the physiological responses of ungulates to dehydration. This overview and annotated bibliography are intended Adaptations for Thermoregulation and as a source of references for wildlife biologists, wildlife managers, and others interested in Maintenance of Water Balance wildlife who have a modest amount of knowledge and experience with animal physiology. The physiological adaptations possessed by ungulates for thermoregulation and maintenance At the outset, it is necessary to define water of water balance have been investigated using a balance and water flux because they commonly variety of methods in domestic goats, sheep, appear in the literature contained in this horses, donkeys (Equus asinus), cattle, camels bibliography. Water balance is the condition (Camelus dromedarius), guanacos (Llama where intake of free water, preformed water in guanicoe), llamas (Llama glama), cervids (deer food, and metabolic water equals water loss [Odocoileus spp.] and elk [Cervus elaphus]), through feces, urine, and cutaneous and pronghorn (Antilocapra americana), desert pulmonary evaporation over a specific period