Antelope Jackrabbit (Lepus alleni) Spatial Ecology, HabitatCharacteristics, and Overlap with the Endangered Pima

Pineapple Cactus (Coryphantha scheeri var. Robustispina)

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Authors Altemus, Maria Michael

Publisher The University of Arizona.

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Download date 28/06/2018 15:03:25

Link to Item http://hdl.handle.net/10150/613565

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ANTELOPE JACKRABBIT (LEPUS ALLENI) SPATIAL ECOLOGY, HABITAT

CHARACTERISTICS, AND OVERLAP WITH THE ENDANGERED PIMA

PINEAPPLE CACTUS (CORYPHANTHA SCHEERI VAR. ROBUSTISPINA)

by

Maria M. Altemus

____________________________ Copyright © Maria M. Altemus 2016

A Thesis Submitted to the Faculty of the

SCHOOL OF NATURAL RESOURCES AND THE ENVIRONMENT

In Partial Fulfillment of the Requirements

For the Degree of

MASTER OF SCIENCE

WITH A MAJOR IN NATURAL RESOURCES

In the Graduate College

THE UNIVERSITY OF ARIZONA

2016

2

STATEMENT BY AUTHOR

The thesis titled Antelope Jackrabbit (Lepus alleni) Spatial Ecology, Habitat

Characteristics, and Overlap with the Endangered Pima Pineapple Cactus

(Coryphantha scheeri var. robustispina) prepared by Maria M. Altemus has been

submitted in partial fulfillment of requirements for a master’s degree at the University

of Arizona and is deposited in the University Library to be made available to borrowers

under rules of the Library.

Brief quotations from this thesis are allowable without special permission,

provided that an accurate acknowledgement of the source is made. Requests for

permission for extended quotation from or reproduction of this manuscript in whole

or in part may be granted by the head of the major department or the Dean of the

Graduate College when in his or her judgment the proposed use of the material is in

the interests of scholarship. In all other instances, however, permission must be

obtained from the author.

SIGNED: _______________________________

Maria M. Altemus

APPROVAL BY THESIS DIRECTOR

This thesis has been approved on the date shown below:

__13 May 2016___

John L. Koprowski Date

Professor of Wildlife and Fisheries Science

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ACKNOWLEDGEMENTS

I would like to thank my major advisor, John L. Koprowski, for his support and

guidance through my graduate program. Thanks to Robert Mannan for helpful

suggestions throughout the project. Special thanks to David E. Brown for sharing his

passion and wisdom of Arizona’s super bunny with me. Sincere thanks to our lab

manager, Vicki L. Greer, and my supportive lab mates (Team KowPow), including

Samuel Abercrombie, Kendell R. Bennett, Hsiang Ling Chen, Jonathan J. Derbridge,

Kirsten Fulgham, Emily A. Goldstein, Sarah L. Hale, Timothy G. Jessen, Shari L.

Ketcham, Allyssa L. Kilanowski, Maxwell N. Mazzella, Calebe P. Mendes, Melissa J.

Merrick, and Amanda M. Veals, who took the time to review and comment on my drafts

and presentations. Thank you to Lacrecia Johnson, who was instrumental in making this

project a reality. A huge shout out to Jim “Jackrabbit Jim” Heffelfinger for his insight on

antelope jackrabbits. Thank you to Randy Babb for his great stories, always interesting

conversations, and guidance. Thanks to Brent Sigafus for his logistical assistance in the

field. Thank you to the staff of Buenos Aires National Wildlife Refuge, including Sally

Flatland, for being supportive of this study and providing PPC location data. This project

would not have been possible without the numerous volunteers and students willing to

run through the desert at night to help try to catch antelope jackrabbits. Without their

assistance, especially Jesse J. Altemus, Nerissa V. Hall, and Cynthia L. Norton, I would

still be chasing antelope jackrabbits. I would like to thank Alberto Moreno for his

permanently sunny attitude, support and love, and assistance with vegetation

measurements. Thank you to my friends for always being there for me, especially

Kimberly Ludwig, Chris Mayer, Melissa K. McRoberts, and Tana Phillips. I cannot

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thank my family enough for their love and encouragement, especially my mom, Gloria V.

Altemus, who always took care of my menagerie whenever I needed her to.

Funding was provided by the Reid Park Zoological Society, T&E Inc., the Desert

Southwest Cooperative Ecosystem Studies Unit (DSCESU) in cooperation with the U.S.

Fish and Wildlife Service and National Park Service, Arizona Agricultural Experiment

Station, University of Arizona Vice President for Research, Mt. Graham Red Squirrel

Research Program, Koprowski Conservation Research Laboratory, the Clifford W.

Carstens, Jr. Endowment Scholarship, the Arrington Memorial Scholarship, the

University of Arizona College of Agriculture and Life Sciences, and the American

Society of Mammalogists.

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DEDICATION

For my dad.

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TABLE OF CONTENTS

LIST OF FIGURES……………………………………………...……………………...7

LIST OF TABLES……………………………………………………………………....8

ABSTRACT…………………………………………………………………………....10

CHAPTER 1: INTRODUCTION…………………………………………………….....11

Animal Home Range…………………………………………………………….11

Vegetation Selection………….………………………………………………….11

An Endangered Cactus…………………………………..………………………11

Study Organism………………………………………...………………………..12

Literature Cited………………………………………………...………………..13

CHAPTER 2: PRESENT STUDY………………………………………………………18

Summary of Methods…………………………………………………….………18

Summary of Results………………………………………………………...……19

Summary of Conclusions…………………......……………………………….....20

APPENDIX A: Seasonal movement patterns and home range of the antelope jackrabbit

(Lepus alleni)………………………….…………………………………...…………….21

APPENDIX B: Antelope jackrabbit (Lepus alleni) habitat characteristics and spatial

association with the Pima pineapple cactus (Coryphantha scheeri var. robustispina) in an

altered semidesert grassland ……………………………………………...……………..49

APPENDIX C: Supplemental data on antelope jackrabbit (Lepus alleni) captures, core

areas, home ranges, and seasonal ranges ………………………………..………………80

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LIST OF FIGURES

FIGURE A1. Core area and home range estimates for 7 antelope jackrabbits (Lepus

alleni) collected from December 2013–November 2015 on Buenos Aires National

Wildlife Refuge, Arizona…………………………………………………………..….....46

FIGURE A2. Core area and home range estimates for 5 antelope jackrabbits (Lepus

alleni) collected from December 2013–November 2015 on Buenos Aires National

Wildlife Refuge, Arizona……………………………..………………………………....47

FIGURE A3. Mean (± SD) of core area and home range estimates (ha) of 12 antelope

jackrabbits (Lepus alleni) collected from December 2013–November 2015 on Buenos

Aires National Wildlife Refuge, Arizona …………………………………………….....48

FIGURE B1. Mean ± SE vegetation obstruction measurements (VOMs) in cm at antelope

jackrabbit (Lepus alleni) use and available vegetation points on Buenos Aires National

Wildlife Refuge, Arizona taken between October–December 2015....………….……....77

FIGURE B2. Differences between the average percentages of each dominant vegetation

characteristic at used vegetation points and available vegetation points on Buenos Aires

National Wildlife Refuge, Arizona for antelope jackrabbit (Lepus alleni). Abbreviations

are amaranth (AMAR), bare ground (BARE), cactus (CACT), desert broom (DEBR),

forbs (FORB), native grass species (GRAS), Johnson grass (JOHN), Lehmann lovegrass

(LEHM), mesquite species (MESQ), palo verde (PALV), snakeweed (SKWD), and

wolfberry (WOLF). Measurements were taken between taken between October–

December 2015…………………………………………………………..………….....78

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LIST OF TABLES

TABLE A1. Home range estimates of various hare species gleaned from the literature..44

TABLE A2. Mean ± SD (ha) antelope jackrabbit (Lepus alleni) seasonal core (50% fixed

kernel) and seasonal range (95% fixed kernel) density estimates and mean home range

usage percentage from December 2013–November 2015 on Buenos Aires National

Wildlife Refuge, Arizona…………..………………..………………………….………..45

TABLE B1. Mean percentage of each dominant vegetation group at use points in

antelope jackrabbit (Lepus alleni) habitat and at available points for all locations on

Buenos Aires National Wildlife Refuge, Arizona, measured October–December 2015...79

TABLE C1. Capture history for all antelope jackrabbits caught on Buenos Aires National

Wildlife Refuge, Arizona, between December 2013–November 2015. Fates are of

animals collared and indicate whether the collars of animals were found (mortality),

whether the collars stopped transmitting a signal and therefore we could not determine a

fate (missing), or whether the animal’s collar was still transmitting a signal at the end of

the study (alive)…………………………………………………………………………..81

TABLE C2. Number of antelope jackrabbits (Lepus alleni) caught and collared, by sex,

on Buenos Aires National Wildlife Refuge, Arizona, between December 2013–November

2015………………………………………………………………………………………83

TABLE C3. Number of antelope jackrabbits (Lepus alleni) caught and collared, by age

group, on Buenos Aires National Wildlife Refuge, Arizona, between December 2013–

November 2015. ………………………………………………………………………....83

TABLE C4. Core area (50% fixed kernel), home range (95% fixed kernel) and seasonal

range (95% fixed kernel) density estimates (ha) for individual antelope jackrabbits (Lepus

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alleni) between December 2013–November 2015 on Buenos Aires National Wildlife

Refuge, Arizona. Winter months were December, January, February and March. Spring

months were April, May, and June. Monsoon months were July, August, and September.

Fall months were October and November…...…………………………..………………84

TABLE C5. Home range usage percentage for each season and degree of interseason

similarity (DIS) for all seasons for each antelope jackrabbit (Lepus alleni) from

December 2013–November 2015 on Buenos Aires National Wildlife Refuge, Arizona.

Home range usage percentage is the seasonal home range size is divided by the home

range. Lower DIS values indicate a more sedentary lifestyle, whereas higher DIS values

are indicative of actively moving individuals. Winter months were December, January,

February and March. Spring months were April, May, and June. Monsoon months were

July, August, and September. Fall months were October and November…………...….85

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ABSTRACT

The antelope jackrabbit (Lepus alleni) inhabits the seasonal landscape of the subtropical

Sonoran savanna grassland in southern Arizona. Basic ecological information on this

understudied lagomorph is lacking beyond historical responses to rangeland conditions.

This is the first study to utilize radio collars to assess space use of antelope jackrabbits.

In the semidesert grassland of Buenos Aires National Wildlife Refuge, Arizona, we

estimated antelope jackrabbit home range size, seasonal ranges, and movement patterns.

Home range estimates were comparable to other Lepus species, however, seasonal range

sizes did not differ. We analyzed antelope jackrabbit habitat structure, measured

vegetation characteristics, and determined whether there was a spatial association

between antelope jackrabbits and the endangered Pima pineapple cactus (Coryphantha

scheeri var. robustispina). Antelope jackrabbits selected vegetation structure and

characteristics similarly to available areas on the refuge. We did not detect a spatial

association between antelope jackrabbits and Pima pineapple cacti, however given the

importance of understanding endangered species relationships, further investigation is

warranted. Our results add to the limited ecological information known about antelope

jackrabbits and provide baseline data for future studies. Knowledge about spatial

ecology and habitat selection helps managers and biologists make informed

recommendations for land and wildlife management.

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CHAPTER 1: INTRODUCTION

Animal Home Range

The home range of an animal is the area that an animal maintains throughout life

while searching for food, mates, and raising young (Burt 1943). Studying the home

range dynamics of many individuals provides a wealth of information regarding the

social and mating systems of that species (Greenwood 1980; Damuth 1981; Parker and

Waite 1997; Cooper and Randall 2007). Additionally, home range size of mammals,

including lagomorphs, is often a reflection of resource quality or availability (e.g., Hixon

1980; Powers and McKee 1994; Powell 2000).

Vegetation Selection

Jackrabbits require heterogeneity in vegetation height and species diversity to

provide food resources and enhance detection and evasion of predators (Lechleitner

1958; Longland 1991; Marín et al. 2003; Smith et al. 2004). For some bird species,

abundance and density in areas with native and non-native grasses do not differ,

indicating that some species place more importance on structure rather than composition

(e.g., Lloyd and Martin 2005; Flanders et al. 2006; Kennedy et al. 2009; Thompson et al.,

2009).

An Endangered Cactus

The Pima pineapple cactus (Coryphantha scheeri var. robustispina; PPC) is a

small cactus, measuring 10–46 cm tall and 7.5–18 cm in diameter, found in Pima and

Santa Cruz counties in southern Arizona and northern Sonora, Mexico (Benson 1982;

Arizona Ecological Services Field Office 2000). It was listed as endangered in the

United States under the Endangered Species Act in 1993. PPC inhabit gently rolling

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alluvial fans within desert grasslands and desert scrubland (Benson 1969; Roller 1996;

McDonald and McPherson 2005; McDonald 2007; USFWS 2007). Factors that

influenced the listing of the PPC in the US included habitat modification and degradation,

illegal collection, disease, predation, and lack of legal protection (USFWS 1993).

Although the PPC is listed as endangered, our knowledge of its basic ecology and

interaction with pollinators and seed dispersers is limited. Rabbits and jackrabbits are

seed dispersers for various plant species, including cactus (Riegel 1941; Cook 1942;

Timmons 1942; Zedler and Black 1992) and rodent and lagomorph pellets are often

found near PPC during the fruiting season (Roller 1996).

Study Organism

The antelope jackrabbit (Lepus alleni) inhabits the tropic-subtropic Sonoran

savanna grasslands of southern Arizona and is the second largest hare in North America

(Nelson 1909; Townsend 1912). Antelope jackrabbit distribution stretches from southern

Arizona south to the Mexican states of Sonora, Sinaloa, and Nayarit (Nelson 1909;

Townsend 1912). Adult antelope jackrabbit weight ranges between 2.7–5.9 kg (Vorhies

and Taylor 1933). The antelope jackrabbit does not exhibit sexual dimorphism, so it is

not possible to differentiate the sexes at a distance (Flux and Angermann 1990). Both

males and females are gray and brown in color, and flash their large white flanks when

alarmed (Hoffmeister 1986). The white-lined ears of the antelope jackrabbit are a

striking identifier, with a length of 162 mm (Vorhies and Taylor 1933). The antelope

jackrabbit is mostly inactive during the day, choosing to hide at the base of a mesquite

tree or under a bush in a small indentation, known as a form, for shelter and to

behaviorally thermoregulate. Antelope jackrabbits leave the form in the late afternoon

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hours and will remain active throughout the night into midmorning. The antelope

jackrabbit’s diet consists of, in order of consumption, grasses, mesquite, cactus, and other

miscellaneous vegetation (Vorhies and Taylor 1933). The percentage of each plant group

consumed varies depending on season and food availability (Vorhies and Taylor 1933).

Grassy hills at elevations from sea level to 1912 m. are preferred, but antelope jackrabbits

will occupy a range of habitats, including the bajada above cactus belts, creosote desert,

and mesquite-laden basins (Vorhies and Taylor 1933).

Literature Cited

Arizona Ecological Services Field Office. 2000. Pima pineapple cactus (Coryphantha

scheeri var. robustispina).

<http://www.fws.gov/southwest/es/arizona/Documents/Redbook/Pima%20Pineap

ple%20cactus%20RB.pdf>. Accessed 22 Jan 2016.

Benson, L. 1969. The cacti of Arizona. Third edition. University of Arizona Press,

Tucson, AZ, USA.

Benson, L. 1982. The cacti of the United States and Canada. Stanford University Press,

Stanford, CA, USA.

Burt, W. H. 1943. Territoriality and home range concepts as applied to mammals. Journal

of Mammalogy 24: 346–352.

Cook, C. W. 1942. Insects and weather as they influence growth of cactus on the Central

Great Plains. Ecology 23: 209–214.

Cooper, L. D., and J. A. Randall. 2007. Seasonal changes in home ranges of the giant

kangaroo rat (Dipodomys ingens): a study of flexible social structure. Journal of

Mammalogy 88: 1000–1008.

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Damuth, J. 1981. Home range, home range overlap, and species energy use among

herbivorous mammals. Biological Journal of the Linnean Society 15: 185–193.

Flanders, A. A., W. P. Kuvlesky, Jr., D. C. Ruthven, III, R. E. Zaiglin, R. L. Bingham, T.

E. Fulbright, F. Hernandez, and L. A. Brennan. 2006. Effects of invasive exotic

grasses on South Texas rangeland breeding birds. Auk 123: 171–182.

Flux, J. E. C., and R. Angermann. 1990. Chapter 4: the hares and jackrabbits. Pages 61–

94 in J. A. Chapman, and J. E. C. Flux, editors. Rabbits, hares, and pikas: status

survey and conservation action plan. IUCN, Gland, Switzerland.

Greenwood, P. J. 1980. Mating systems, philopatry, and dispersal in birds and mammals.

Animal Behaviour 28: 1140–1162.

Hixon, M. A. 1980. Food production and competitor density as the determinants of

feeding territory size. American Naturalist 122: 366–391.

Hoffmeister, D. F. 1986. Mammals of Arizona. The University of Arizona Press, Tucson,

AZ, USA.

Kennedy, P. L., S. J. DeBano, A. M. Bartuszevige, and A. S. Lueders. 2009. Effects of

native and exotic grassland plant communities on breeding passerine birds:

implications for restoration of northwest bunchgrass prairie. Restoration Ecology

17: 515–525.

Lechleitner, R. R. 1958. Movements, density, and mortality in a black-tailed jack rabbit

population. The Journal of Wildlife Management 22: 371–384.

Lloyd, J. D., and T. E. Martin. 2005. Reproductive success of chestnut-collared longspurs

in native and exotic grassland. The Condor 107: 363–374.

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Longland, W. S. 1991. Risk of predation and food consumption by black-tailed

jackrabbits. Journal of Rangeland Management 44: 447–450.

Marín A. I., L. Hernández, and J. W. Laundré. 2003. Predation risk and food quantity in

the selection of habitat by black-tailed jackrabbit (Lepus californicus); and

optimal foraging approach. Journal of Arid Environments 55: 101–110.

McDonald, C. J. 2007. Pima pineapple cactus: a unique cactus hiding in plain sight. The

Plant Press 31: 1–4.

McDonald, C., and G. McPherson. 2005. Pollination of Pima pineapple cactus

(Coryphantha scheeri var. robustispina): does pollen flow limit abundance of this

endangered species? Pages 529–532 in Proceedings: Connecting mountain islands

and desert seas: biodiversity and management of the Madrean Archipelago II

symposium. USDA Forest Service, Tucson, AZ, USA.

Nelson, E. W. 1909. The rabbits of North America. North American Fauna 29: 1–314.

Parker, P. G., and T. A. Waite. 1997. Mating systems, effective population size, and

conservation of natural populations. Pages 243–261 in J. R. Clemmons, and R.

Buchholz, editors. Behavioral approaches to conservation in the wild Cambridge

University Press, Cambridge, United Kingdom.

Powell, R. A. 2000. Home ranges, territories, and home range estimators. Pages 65–110

in L. Boitani, and T. Fuller, editors. Research techniques in animal ecology:

controversies and consequences. Columbia University Press, New York, NY,

USA.

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Powers, D. R., and T. McKee. 1994. The effect of food availability on time and energy

expenditure of territorial and non-territorial hummingbirds. Condor 96: 1064–

1075.

Riegel, A. 1941. Some coactions of rabbits and rodents with cactus. Transactions of the

Kansas Academy of Science 44: 96–101.

Roller, P. S. 1996. Distribution, growth, and reproduction of Pima pineapple cactus

(Coryphantha scheeri Kuntz var. robustispina Schott) [M. S. Thesis] University

of Arizona, Tucson, AZ, USA.

Smith R. K., N. V. Jennings, A. Robinson, and S. Harris. 2004. Conservation of

European hares (Lepus europaeus) in Britain: is increasing habitat heterogeneity

in farmland the answer? Journal of Applied Ecology 41: 1092–1102.

Thompson, T. R., C. W. Boal, and D. Lucia. 2009. Grassland bird associations with

introduced and native grass Conservation Reserve Program fields in the Southern

High Plains. Western North American Naturalist 69: 481–490.

Timmons, F. L. 1942. Dissemination of prickly pear seed by jack rabbits. Journal of the

American Society of Agronomy 34: 513–520.

Townsend, C. H. 1912. Mammals collected by the 'Albatross' expedition in Lower

California in 1911, with descriptions of new species. Bulletin of the American

Museum of Natural History 31: 117–130.

U. S. Fish and Wildlife Service [USFWS]. 1993. Endangered and threatened wildlife and

plants; determination of endangered status for the plant Pima pineapple cactus

(Coryphantha scheeri var. robustispina). Federal Register Vol. 58 No. 183, 50

CFR Part 17, RIN 1018-AB75.

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<http://www.fws.gov/southwest/es/arizona/Documents/SpeciesDocs/PimaPineapp

leCactus/Pima%20Pineapple%20Cactus.pdf>. Accessed 22 Jan 2016.

U.S. Fish and Wildlife Service [USFWS]. 2007. U.S. Fish & Wildlife Service 5-year

review determination Pima pineapple cactus. 70 FR 5460. Region 2 Regional

Office.

Vorhies, C. T., and W. P. Taylor. 1933. The life histories and ecology of jack

rabbits, Lepus alleni, and Lepus californicus ssp., in relation to grazing in

Arizona. Arizona Agricultural Experiment Station Technical Bulletin 49: 471–

587.

Zedler, P. H., and C. Black. 1992. Seed dispersal by a generalized herbivore: rabbits as

dispersal vectors in a semiarid California vernal pool landscape. The American

Midland Naturalist 128: 1–10.

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CHAPTER 2: PRESENT STUDY

This thesis is comprised of two manuscripts. These manuscripts can be found in

Appendices A and B. Appendix A, formatted for the Journal of Mammalogy, entitled,

“Seasonal movement patterns and home range of the antelope jackrabbit (Lepus alleni)”,

assesses home range size and seasonal movement patterns of antelope jackrabbits (Lepus

alleni). Appendix B, formatted for Rangeland Ecology and Management, entitled,

“Antelope jackrabbit (Lepus alleni) habitat characteristics and spatial association with the

Pima pineapple cactus (Coryphantha scheeri var. robustispina) in an altered semidesert

grassland”, examines antelope jackrabbit habitat characteristics and the spatial

association with the Pima pineapple cactus (Coryphantha scheeri var. robustispina). The

following is a summary of the major findings.

Summary of Methods

This study occurred on the Buenos Aires National Wildlife Refuge in Pima

County in southcentral Arizona, USA. From December 2013–July 2015 we captured

adult and juvenile antelope jackrabbits, ear-tagged all animals, collected biological data,

and radio-collared adults. We tracked 13 animals from December 2013–November 2015

and calculated 95% fixed kernel (home range) and 50% fixed kernel (core area) density

estimates; we also calculated seasonal ranges for four biologically significant seasons:

monsoon (wet), fall (dry), winter (wet), and spring (dry).

We established “use” vegetation points in antelope jackrabbit home ranges and

“available” vegetation points within the study area to determine habitat structure and

characteristics. We measured visual obstruction measurements (VOM) and various

vegetation characteristics at each vegetation point; we pooled individual vegetation

19

characteristics into vegetation groups. We compared distance of antelope jackrabbits

telemetry locations and random points to Pima pineapple cactus locations to determine if

a spatial association existed.

Summary of Results

Mean ± SD core area was 5.20 ± 3.87 ha and mean home range was 29.14 ± 15.58

ha (n = 12). Male and female home range size did not differ. Mean ± SD seasonal range

size varied from 15.44 ± 10.78 ha in the fall to 29.09 ± 23.98 ha in the spring. Seasonal

range sizes did not differ, although seasonal ranges shifted slightly from season to season.

Mean ± SD VOM for areas of use was 12.68 ± 8.87 cm, whereas mean VOM for

available points was 13.04 ± 13.57 cm. VOMs between all use and available vegetation

points did not differ and vegetation groups did not differ between all use and available

points. Percentages of antelope jackrabbit points and random points near Pima pineapple

cacti did not differ, however distances of antelope jackrabbit points to Pima pineapple

cactus locations were larger than distances of random points to Pima pineapple cacti.

During the fruiting season, between August and December, antelope jackrabbit locations

were not closer to Pima pineapple cactus than random points to Pima pineapple cactus.

Summary of Conclusions

Antelope jackrabbit home range estimates were comparable to other Lepus

species. The lack of seasonal range differences in antelope jackrabbits is surprising given

the seasonality of the landscape and reports of seasonal ranges in other jackrabbits.

Vegetation structure and composition in antelope jackrabbit home ranges were similar to

that available; vegetation obstruction measurements did not predict home range size,

indicating other factors besides vegetation height and species are influencing habitat

20

selection. Our results add to the limited ecological information known about antelope

jackrabbits and provide a basis for future studies, and will help land managers create

recommendations for land and wildlife management.

21

APPENDIX A: Seasonal movement patterns and home range of the antelope jackrabbit

(Lepus alleni)

(Formatted for Journal of Mammalogy)

Maria M. Altemus, School of Natural Resources and the Environment, The University of

Arizona, 1064 E. Lowell St., Tucson, AZ 85721, maltemus@email.arizona.edu

Antelope jackrabbit space use

Seasonal movement patterns and home range of the antelope jackrabbit (Lepus

alleni)

Maria M. Altemus*, John L. Koprowski, and David E. Brown

School of Natural Resources and the Environment, University of Arizona, Tucson, AZ

85721 USA (MMA, JLK); School of Life Sciences, Arizona State University, Tempe, AZ

85281 USA (DEB)

In the face of climate change, more research is needed to understand how animals adjust

to dynamic ecosystems. Seasonal environments offer a unique opportunity to observe

how animals use the landscape in a fluctuating system. The antelope jackrabbit (Lepus

alleni) is an understudied lagomorph found in the Sonoran Desert of Arizona, USA and

Mexico. Basic ecological information on this species is lacking beyond historical

responses to rangeland conditions. We used radio telemetry to collect novel data on

home range size and seasonal movement patterns of antelope jackrabbits on the Buenos

Aires National Wildlife Refuge in southcentral Arizona. Male and female antelope

jackrabbit mean home range size did not differ. Despite the seasonal landscape of

semidesert grasslands, antelope jackrabbits home range size did not change across

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seasons. Antelope jackrabbit home ranges are comparable to range sizes reported for the

widespread black-tailed jackrabbit (Lepus californicus) as well as the endangered

Tehuantepec jackrabbit (Lepus flavigularis). This study is the first to document the home

range of antelope jackrabbits, and our results will allow us to better understand how hares

are using the landscape in a complex and ever-changing arid environment.

Key words: Arizona, semidesert grassland, hare, lagomorph, Sonoran Desert, space use,

spatial ecology

*Correspondent: maltemus@email.arizona.edu

23

The Sonoran Desert is North America’s hottest desert and is already being

affected by climate change, with warmer temperatures and fewer freeze events in recent

times (Weiss and Overpeck 2005). Currently, the Desert Southwest of southern Arizona,

USA, experiences drawn-out winter rains, followed by dry, hot, summers, punctuated by

intense late-summer monsoons (Arizona-Sonora Desert Museum 2000). Native wildlife

species are well-adapted for the extreme conditions of desert ecosystems (Evenari et al.

1971; Stafford Smith and Morton 1990; Morton et al. 2011). Appropriate responses by

wildlife to cyclic changes in resources include altering dietary choices (Hofmann 1989)

or seasonal variations in home range size (Burt 1943).

A fundamental aspect of species ecology in a seasonal landscape is the temporal

stability of the individual home range. An animal’s home range is the area that an animal

maintains throughout life while searching for food, mates, and raising young (Burt 1943)

and within an individual home range, the core area is considered the space where most

activities are focused (Samuel et al. 1985; Samuel and Garton 1987). Home range is

determined using observational studies and radio telemetry, and more recently, GPS

technology (MacDonald et al. 1980; Millspaugh and Marzluff 2001; Cagnacci et al.

2010). Home range dynamics of multiple individuals provides a wealth of information

regarding the social and mating systems for that species (Greenwood 1980; Damuth

1981; Parker and Waite 1997; Cooper and Randall 2007). Additionally, home range size

of mammals, including lagomorphs, is often a reflection of resource quality (e.g., Hixon

1980; Powers and McKee 1994; Powell 2000).

24

Examining how animals use the landscape not only improves our knowledge of

species’ ecology, but can also inform us about resource availability and habitat quality

(Ford 1983). For example, hare home range size varies due to differences in habitat

cover, resources, and water availability (Lechleitner 1958; Smith 1990; Flinders and

Chapman 2003). Among Lepus species, home range sizes vary intraspecifically

depending on location (Table 1; Kunst et al. 2001; Stott 2003; Rühe and Hohmann 2004).

Variation in resource availability and quality has likely led to a wide range of home range

estimates in the black-tailed jackrabbit (Table 1). In highly seasonable environments,

animals shift home ranges and make movements in response to fluctuating environmental

conditions (Ferguson et al. 1999). For example, the grassland desert-inhabiting

Tehuantepec jackrabbit (Lepus flavigularis) shows seasonal variation in home range size

(Carrillo-Reyes et al. 2010). Home range estimates and seasonal movement patterns for

the desert-dwelling antelope jackrabbit (Lepus alleni) have not been reported.

The antelope jackrabbit is a conspicuous, but understudied Sonoran Desert

mammal (Hoffmeister 1986). Current knowledge of the species is limited to life history

characteristics in relation to cattle grazing in southern Arizona (Vorhies and Taylor

1933), populations dynamics in response to precipitation (Madsen 1974; Gray 1977),

forage consumption (Arnold 1942), and physiology (Dawson and Schmidt-Nielsen 1966;

Hinds 1977). Data on basic ecological traits like home range size, seasonal landscape

use, and habitat preferences have not been fully explored.

Interest in the antelope jackrabbit’s role in the ecosystem has been revitalized by

recent literature reviews and observational surveys (Brown et al. 2014; Lorenzo et al.

2014), but systematic field studies are still lacking. Our study is the first to examine

25

antelope jackrabbit spatial ecology in the semidesert grassland of the Sonoran Desert.

Our objectives were to determine home range size and determine whether antelope

jackrabbits respond to the seasonably variable environment in an expected manner.

Given the seasonality of the Desert Southwest, we would expect antelope jackrabbit

home range to fluctuate based upon resource availability and quality.

Methods

Study Site and Study Animal.—This study was conducted on the Buenos Aires National

Wildlife Refuge (31.5501° N, 111.5507° W; BANWR), in Pima County in southcentral

Arizona, 90 km southwest of Tucson, Arizona. BANWR encompasses 46,450 ha in the

Altar Valley basin. The U.S.-Mexico international border is the southern boundary of

BANWR, with the Baboquivari Mountains to the west and the Sierrita Mountains to the

east. Elevation ranges between 925 to 1,400 m (Koenen and Krausman 2002), however

most of the refuge occurs within 950 to 1,150 m (McLaughlin 1992). BANWR is

comprised mainly of a semidesert grassland and Sonoran desertscrub, with relict

subtropical savanna grassland (Brown 1994). Exotic grasses (Eragrostis chloromelas, E.

lehmanniana, Sorghum halepense), shrubs (Ambrosia dumosa, Baccharis sarothroides,

Gutierrezia sarothrae), native grasses (Aristida hamulosa, A. ternipes, Bouteloua

rothrocku, B. repens, Digitaria californica), a variety of cactus species (Carnegiea

gigantea, Echinocereus fasciculatus, Opuntia spp.) and velvet mesquite (Prosopis

velutina) stands can be found on the refuge (McLaughlin 1992; Brown 1994). BANWR

receives an average of 410 mm annual precipitation (Sellers et al. 1985), with monsoon

rains occurring in July and August and winter rains occurring between December and

March (Arizona-Sonora Desert Museum 2000; McLaughlin 1992).

26

Capture.— We traversed unpaved roads of BANWR at night and used a spotlight to

locate antelope jackrabbits (FatMax 900 Lumen LED, Stanley Tools, Towson, MD). We

used a hoop net (Mighty Net 34”x40”, TJB Inc., Hamden, CT) or handheld net launcher

(MagNet, Wildlife Capture Services, Flagstaff, AZ) to capture animals . After capture,

we transferred animals to a black nylon sack (approximately 71 cm by 96 cm) for

handling. To weigh animals, we used a 30 kg capacity hanging scale (UltraSport V2-30,

JScale, Phoenix, AZ) and measured the right hind foot. We documented the sex and

reproductive status of each animal. We used a self-piercing metal numbered ear tag

(Monel #3, National Band & Tag, Newport, KY) to ear-tag each animal. We fitted VHF

radio collars (Model LPM-2180A, Wildlife Materials, Murphysboro, IL) on adult animals

and released them at the site of capture. All field work was conducted in accordance with

the American Society of Mammalogists guidelines and approved by the University of

Arizona’s Institutional Animal Care and Use Committee (Sikes et al. 2011; IACUC

protocol #13-480). Field methods were conducted under permits from the Arizona Game

and Fish Department (LIC #SP654189) and Buenos Aires National Wildlife Refuge

(Special Use Permit #2013-028).

Home Range Size.—We located animals at least once per week, between 0800-0030

hours, with a 3-element folding Yagi antenna (Wildlife Materials, Inc., Murphysboro, IL)

and VHF receiver (TRX-2000S, Wildlife Materials, Inc., Murphysburo, IL). We used

single-person biangulation to determine animal location, with interbearing intervals < 10

min. of one another. We used Backpacker GPS Trails (Trimble Navigation Ltd.,

Sunnyvale, CA to record animal locations.

27

We used the “best biangulation” function in Location of a Signal (LOAS;

Ecological Software Solutions LLC, Hegymagas, Hungary) to estimate animal locations.

We ran the MCP Sample Size Bootstrap analysis in ArcView (3.3, Environmental

Systems Research Institute Inc. (ESRI), Redlands, CA) as a function of animal locations

to determine the asymptote for home range size. We used the Spatial Analyst (ESRI) and

Animal Movement extensions (Hooge and Eichenlaub 1997; Laver and Kelly 2008) in

ArcView to calculate 95% (home range) and 50% (core area) fixed kernel density

estimates. We performed t-tests to compare core area and home range sizes between

females and males.

Seasonal Range Size and Movement.— We separated animal locations into four

biologically relevant seasons: winter (wet), spring (dry), monsoon (wet), and fall (dry).

Winter months were December, January, February and March. Spring months were

April, May, and June. Monsoon months were July, August, and September. Fall months

were October and November. We estimated 50% fixed kernel density estimates

(“seasonal core”) and 95% fixed kernel density estimates (“seasonal range”) for each

animal in each season. We calculated seasonal ranges for animals that had locations in

each season.

We used a repeated measures analysis of variance test (ANOVA) to compare

mean seasonal core size and seasonal range size among seasons (Keppel 1991). We used

a multiple regression analysis to determine the effect of sex, season, and site on the

seasonal range size.

For each animal, we calculated the area (ha) that each seasonal range overlapped

with the home range; we took that area and divided it by the home range to calculate

28

home range (HR) usage percentage. For each season, we averaged all HR usage

percentages. We compared mean HR usage percentages between seasons using a single

factor ANOVA (Sokal and Rohlf 1981).

For each animal, we calculated the area (ha) that all seasonal ranges overlapped

by using the intersect tool in ArcMap (10.3.1, ESRI) and creating a new polygon (Ov

below). We determined the degree of interseason similarity for each animal

(Misiorowska 2013). The degree of interseason similarity formula is:

DIS =

4Ov

.100[%]

(Wn + Sp + Mn + Fl)

where DIS is the degree of interseason similarity; Ov is the area (ha) that all seasons

overlap; and Wn, Sp, Mn, and Fl are the seasonal range estimates (ha). We multiplied Ov

by four because we have four seasons (Misiorowska 2013). We conducted a t-test to

determine whether a difference existed in degree of interseason similarity between males

and females. Mean ± 1 SD is provided within the results.

Results

Home Range Size.—Between December 2013 and November 2015, we monitored 22

radio-collared animals. We obtained enough locations on 13 animals to calculate home

range estimates. Home range size became stable with between 20–25 locations; we

calculated home range estimates for animals (n = 13) with at least 23 locations, averaging

of 47.8 ± 23.2 locations per animal (Figures 1 and 2). We calculated home range

estimates for 6 females and 7 males, however one female (F19) had an

uncharacteristically large home range (> 2200% larger than the mean) and was excluded

from the analysis. Mean core area for all animals was 5.20 ± 3.87 ha and mean home

29

range was 29.14 ± 15.58 ha (n = 12) (Figure 3). Core and home range areas tended to be

slightly larger for males (core: t10 = 1.79, P = 0.10; home range: t10 = 2.04, P = 0.07).

Seasonal Range Size and Movement.— We calculated seasonal core area and seasonal

range estimates for 8 animals (5 males, 3 females), collected between December 2013

and November 2015. Mean number of locations per animal per season was 14.69 ±

10.05.

Seasonal core area sizes and seasonal range sizes did not differ for any season

(Table 2; core by season: F3, 21 = 0.59, P = 0.63; range by season: F3, 21 = 2.22, P = 0.12).

The relationship between seasonal range and site, season, and sex was not linear (F3, 28 =

1.66, P = 0.20, R2 = 0.15), however sex did have significant explanatory power (β =

13.08, t28 = 2.15, P = 0.04). Mean HR usage percentage differed between seasons with

overlap maximal during the monsoon season (Table 2; F3, 28 = 5.01, P < 0.01). Degree of

interseason similarity ranged from 14.97% to 48.72%, with a mean of 27.29% ± 11.46.

The sexes did not differ in degree of interseason similarity (t6 = 1.32, P = 0.23).

Discussion

Home Range Size.—Estimates of antelope jackrabbit home range size were similar to

home range sizes for other Lepus species, including black-tailed jackrabbits (French et al.

1965; Smith 1990), Tehuantepec jackrabbits (Carrillo-Reyes et al. 2010) and European

hares (Table 1; Kunst et al. 2001). Male antelope jackrabbits tended to have larger home

ranges than females, which is similar to black-tailed jackrabbits (Smith 1990) and

Tehuantepec jackrabbits (Farías et al. 2006).

Seasonal Range Size and Movement.— Seasonal range sizes did not differ despite the

seasonality of BANWR and seasonal ranges found in other jackrabbits (Carrillo-Reyes et

30

al. 2010). This is surprising since other lagomorphs, including the mountain hare (Lepus

timidus), have smaller ranges when available biomass was high (Hulbert et al. 1996).

Likewise, black-tailed jackrabbits in Utah had different range sizes through different

seasons, with spring and fall ranges being the smallest (Smith 1990).

Although seasonal range sizes remained stable, differences in HR usage

percentage indicate that antelope jackrabbits shift ranges seasonally. Monsoon (wet)

ranges had the highest HR usage percentage and were only slightly larger than spring

(dry) seasons. Monsoon and spring values were much higher than both winter (wet) and

fall (dry) HR usage percentages. In the desert environment, vegetation quality is

influenced by water availability (Nicholson et al. 1990). However, patterns of antelope

jackrabbit space use did not follow seasonal moisture trends. Animals respond to scarcity

of resources differently. For example, the European hare (Lepus europaeus) restricts

movements to save energy (Kunst et al. 2001), whereas mountain hares have larger home

range sizes when food availability is low (Hulbert et al. 1996). Alternatively, longer

range movements leading to shifting seasonal ranges could be a result of other influences

including breeding dispersal or exploration (Baars 1979; Greenwood 1980; Van Dyck

and Baguette 2005). Because jackrabbit breeding occurs year-round, animals may move

across the landscape in search of mates constantly (Vorhies and Taylor 1933; Madsen

1974).

Mean degree of interseason similarity was relatively low, indicating animals

move throughout home ranges widely, also evidenced by differences in HR usage

percentages. Lower degree of interseason similarity indicates that animals may not be

31

using the same areas for different seasonal ranges, whereas, a higher value is indicative of

an inactive and sedentary lifestyle (Misiorowska 2013).

Studies on Lagomorphs.—Research on understudied species is important for conservation

and management purposes, as well as for understanding a species’ role in overall

ecosystem function. While the ecology and behavior of some lagomorphs is well-

studied, many less common and endemic species have not been thoroughly investigated

(Chapman and Flux 2008). Further knowledge of complex ecological relationships, such

as landscape use and habitat selection, are critical to provide insight into how lagomorph

populations will respond to future challenges such as climate change (Beever et al. 2003;

Morrison and Hik 2007).

Climate Change.—The southwestern United States will be disproportionately affected by

climate change in drastic ways because of the already extreme nature of the environment

(Garfin et al. 2014), and climate models foresee conditions becoming hotter and more

arid (Seager et al. 2007; Diffenbaugh et al. 2008; Wise 2009). Changes in vegetation

distribution, severe droughts, warmer temperatures, increased fire intensity, and further

scarcity of water may force animals to shift their ranges (Weiss and Overpeck 2005;

Garfin et al. 2014; IPCC 2014). The antelope jackrabbit inhabits the Sonoran Desert in

Arizona, USA, and Sonora, Mexico, and its current distribution extends further south into

Sinaloa and Nayarit, Mexico. Sinaloa and Nayarit are less arid than the Sonoran Desert

and are composed of thornscrub and tropical deciduous forest (Krizman 1972; Gentry

1995), which may provide a more temperate refuge if the antelope jackrabbit cannot

adapt to future climate change conditions.

32

Current Management Implications.—Access to current data on spatial ecology will allow

managers to make more informed decisions and determine how land management

practices, like prescribed burns, may affect animals and home range dynamics (McGee

1982; Hobbs and Spowart 1984; Burrow et al. 2000; Rühe and Hohmann 2004; Lorenzo

et al. 2008; Amacher et al. 2011). Additionally, future management strategies will require

sound and recent data to allow land and wildlife managers to mitigate the effects of

climate change (Mawdsley et al. 2009). Our study provides baseline data for further

studies to develop hypotheses on the impacts of resources and land use change on

jackrabbits. Our research investigating seasonal landscape movements and home range

size of antelope jackrabbits provides insight on spatial dynamics of an understudied

species, and will allow mangers to make more informed recommendations and will help

ecologists better anticipate the effects of global change on arid semidesert grassland

systems.

Acknowledgments

This project would not have been possible without the numerous volunteers,

especially J. Altemus, N. Hall, and C. Norton, who helped with animal capture and

telemetry. Sincere thanks to J. Derbridge, E. Goldstein, and A. Veals, who took the time

to review, edit and provide comments on drafts. Many thanks to M. Merrick for her

comments and GIS analysis help. Thanks to J. Heffelfinger for his insight on antelope

jackrabbits. Thanks to B. Sigafus for his help on BANWR.

This work was supported by the Reid Park Zoological Society, T&E Inc.,

USFWS, National Park Service (NPS), and Desert Southwest Cooperative Ecosystem

Studies Unit (DSCESU), Arizona Agricultural Experiment Station, University of Arizona

33

Vice President for Research, Mt. Graham Red Squirrel Research Program, Koprowski

Conservation Research Laboratory, and the American Society of Mammalogists.

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Smith, G. W. 1990. Home range and activity patterns of black-tailed jackrabbits. Great

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Stott, P. 2003. Use of space by sympatric European hares (Lepus europaeus) and

European rabbits (Oryctolagus cuniculus) in Australia. Mammalian Biology 68:

317–327.

Tapia, I. 2010. Genetic diversity and connectivity of white-tailed jackrabbit populations

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44

Figures

Table 1. Home range estimates of various hare species gleaned from the literature.

Author Location Species Mean home range size

Feierabend and Kielland 2014 Alaska L. americanus 3–6 ha

Lechleitner 1958 California L. californicus ~18 ha

French et al. 1965 Idaho L. californicus < 20 ha

Smith 1990 Utah L. californicus <100–300 ha

Kunst et al. 2001 Netherlands L. europaeus 28.7 ha

Rühe and Hohmann 2004 Germany L. europaeus 21 ha

Misiorowska 2013 Poland L. europaeus 43 ha

Farías et al. 2006 Mexico L. flavigularis 55 ha

Carrilo-Reyes et al. 2010 Mexico L. flavigularis 21–26 ha

Sánchez-García et al. 2012 Spain L. granatensis 39.6 ha

Kirk and Bathe 1994 Seychelles L. nigricollis 1.39 ha

Dahl and Willebrand 2005 Sweden L. timidus 198 ha

Kauhala et al. 2005 Finland L. timidus 206 ha

Gamboni et al. 2008 Switzerland L. timidus 38 ha

Donoho 1972 Colorado L. townsendii 259 ha

Schaible 2007 South Dakota L. townsendii 61–200 ha

Tapia 2010 Iowa L. townsendii 32–158 ha

45

Table 2. Mean ± SD (ha) antelope jackrabbit (Lepus alleni) seasonal core (50% fixed

kernel) and seasonal range (95% fixed kernel) density estimates and mean home range

usage percentage from December 2013–November 2015 on Buenos Aires National

Wildlife Refuge, Arizona.

Season Months

Mean

seasonal

core area

Mean

seasonal

range

Mean home

range usage

percentage

Winter (Wet) (Dec, Jan, Feb, Mar) 4.02 ± 2.95 19.94 ± 12.91 53.44 ± 21.29

Spring (Dry) (Apr, May, Jun) 4.78 ± 5.28 29.09 ± 23.98 71.11 ± 16.50

Monsoon (Wet) (Jul, Aug, Sep) 5.08 ± 2.56 28.47 ± 16.90 72.03 ± 13.96

Fall (Dry) (Oct, Nov) 3.37 ± 2.89 15.44 ± 10.78 41.02 ± 22.48

46

Figure 1. Core area and home range estimates for 7 antelope jackrabbits (Lepus alleni)

collected from December 2013–November 2015 on Buenos Aires National Wildlife

Refuge, Arizona.

47

Figure 2. Core area and home range estimates for 5 antelope jackrabbits (Lepus alleni)

collected from December 2013–November 2015 on Buenos Aires National Wildlife

Refuge, Arizona.

48

Figure 3. Mean (± SD) of core area and home range estimates (ha) of 12 antelope

jackrabbits (Lepus alleni) collected from December 2013 – November 2015 on Buenos

Aires National Wildlife Refuge, Arizona.

49

APPENDIX B: Antelope Jackrabbit (Lepus alleni) Habitat Characteristics and Spatial

Association with the Pima Pineapple Cactus (Coryphantha scheeri var. robustispina) in

an Altered Semidesert Grassland

(Formatted for Rangeland Ecology and Management)

Antelope Jackrabbit (Lepus alleni) Habitat Characteristics and Spatial Association

with the Pima Pineapple Cactus (Coryphantha scheeri var. robustispina) in an

Altered Semidesert Grassland

MARIA M. ALTEMUSa*

, JOHN L. KOPROWSKIa, DAVID E. BROWN

b, LACRECIA

JOHNSONc

aSchool of Natural Resources and the Environment, University of Arizona, Environment

and Natural Resources 2, 1064 E. Lowell Street, Tucson, AZ 85721 USA

bSchool of Life Sciences, Arizona State University, 427 E. Tyler Mall, Tempe, AZ 85281

USA

cU.S. Fish and Wildlife Service, National Wildlife Refuge System, Division of Biological

Sciences, 12661 E. Broadway Boulevard, Tucson, AZ 85748, USA

*Correspondence: Maria M. Altemus, School of Natural Resources and the Environment,

University of Arizona, Environment and Natural Resources 2, 1064 E. Lowell Street,

Tucson, AZ 85721 USA, Tel: +1 520 256 4748. E-mail address:

maltemus@email.arizona.edu (M.M. Altemus)

50

ABSTRACT

Anthropogenic influences, such as cattle grazing, nonnative vegetation introduction, and

fire suppression, have greatly altered grasslands globally, and some wildlife species have

been negatively impacted. As efforts to restore grasslands to a pre-settlement state

continue, we must understand the habitat requirements of various resident species, and

the ecological relationships that exist with other plants and animals, especially

endangered species. We examined habitat selection at the within-study area level by

characterizing vegetation at use locations within antelope jackrabbit (Lepus alleni

Mearns, 1890) home ranges on Buenos Aires National Wildlife Refuge in south central

Arizona to available locations within the study area. We documented proximity of

antelope jackrabbit locations to Pima pineapple cacti (Coryphantha scheeri Kuntz var.

robustispina Schott) to determine whether antelope jackrabbits have a spatial relationship

with this endangered cactus. Vegetation structure and characteristics were similar

between use and available vegetation points. Antelope jackrabbit locations were not

concentrated around Pima pineapple cactus plants. Home range size is not explained by

vegetation biomass, which may indicate that antelope jackrabbits select habitat based on

other factors besides food quantity. Though we did not find a spatial association between

radio-collared antelope jackrabbits and Pima pineapple cactus locations, a possible

relationship warrants further investigation. Being aware of habitat characteristics will

allow land managers to understand how antelope jackrabbits are using the landscape and

better determine how to make management decisions in an altered and continually-

changing semidesert grassland.

51

KEYWORDS Arizona, Buenos Aires National Wildlife Refuge, endangered species,

habitat, hare, semidesert grassland

52

INTRODUCTION

Landscape change has occurred across the world due to anthropogenic factors,

including timber harvesting (Lindenmayer and Fischer, 2006), conversion to cropland

(Knopf, 1994; Landsberg, 1999; Daily 2001), urbanization (Luck et al., 2004),

infrastructure development (Forman and Alexander, 1998), fire suppression (Baker,

1992), and climate change (Peters and Lovejoy, 1992; Hughes, 2000; Thomas et al.,

2004). Domestic livestock have also altered the environment in numerous ways

(Peterken, 1996; Wang and Hacker, 1997; Hobbs, 2001). These novel animals compact

soil (Bezkorowajynj et al., 1993), alter soil dynamics and indirectly promote growth of

invasive vegetation (Janzen, 1983; Carr et al., 1992; Primack, 2002), and introduce

disease to native species (Bengis et al., 2002). Livestock grazing also disturbs the natural

fire regime which allows shrubs to proliferate and can lead to changes in the biotic

community from a grassland to a shrubland (Mistry, 2000; Van Auken, 2000). Grazing

decreases soil moisture and increases light availability (Stoutjesdijk and Barkman, 1992)

and can be beneficial to plant species that prefer those conditions (e.g., Moog et al., 2002;

Wahlman et al., 2002). Additionally, in Finland, cattle grazing positively impacts some

mesic grassland plant species (Pykälä 2005).

European settlers arrived in the southwestern United States with cattle and caused

long-lasting landscape change (Wagoner, 1952; Schickedanz, 1980; Bahre, 1991; Brugge

and Gerow, 2000). Mesquite trees (Prosopis spp.), shrubs, and nonnative vegetation

have increased in the semidesert grassland of southcentral Arizona (Archer 1994; Bock et

al. 2007). Landscape changes, whether anthropogenic of natural, lead to changes in

habitat condition and quality (Morrison et al., 1998).

53

While land managers restore habitat for individual species, understanding the

habitat requirements of all plants and animals in the environment is important to

determine how management decisions will affect the overall ecosystem (Sinclair et al.,

2006). Additionally, biologists learn about animal distribution and abundance by

determining habitat conditions and requirements (Morrison et al., 1998). Two sympatric

and ecologically-linked species in southcentral Arizona include the Pima pineapple

cactus (Coryphantha scheeri Kuntz var. robustispina Schott) and the antelope jackrabbit

(Lepus alleni Mearns, 1890).

The Pima pineapple cactus (PPC) is a small cactus, measuring 10–46 cm tall and

7.5–18 cm in diameter, found in Pima and Santa Cruz counties in southern Arizona and

northern Sonora, Mexico (Benson, 1982; Arizona Ecological Services Field Office, 2000)

and was listed as endangered in the United States under the Endangered Species Act in

1993. PPC inhabit gently rolling alluvial fans within desert grasslands and desert

scrubland (Benson, 1969; Roller, 1996; McDonald and McPherson, 2005; McDonald,

2007; USFWS, 2007). Factors that influenced the listing of the PPC in the US included

habitat modification and degradation, illegal collection, disease, predation, and lack of

legal protection (USFWS 1993). Although the PPC is listed as endangered, our

knowledge of basic ecology and interactions with pollinators and seed dispersers is

limited.

Rabbits and jackrabbits are seed dispersers for various plants, including cactus

species (Riegel 1941; Cook, 1942; Timmons, 1942; Zedler and Black, 1992). Although

the seed dispersal strategies of the PPC have not been fully studied, rodent and

lagomorph pellets are often found near PPC during the fruiting season (Roller, 1996).

54

Additionally, lagomorphs eat PPC fruits, and may be the main predators of PPC fruits

(WestLand Resources, Inc., 2009, unpublished report). PPC may benefit from a

mutualistic relationship with jackrabbits through seed dispersal, however it is unclear if

jackrabbits are seeking out these small endangered cacti as a preferred food source. The

documentation of a spatial relationship between antelope jackrabbits and PPC may

influence future management decisions of both species.

The antelope jackrabbit is an understudied hare found in southern Arizona and

Mexico. Jackrabbits require diverse vegetation heights and species for food resources

and cover from predators (Lechleitner 1958; Longland 1991; Marín et al. 2003; Smith et

al. 2004) and antelope jackrabbits occupy a variety of habitats, including mesas or

bajadas, creosote-filled desert, mesquite bosques, desert scrub, savannas, and dense

scrub; however, tropic-subtropic savannas are the principal habitat (Vorhies and Taylor,

1933; Woolsey, 1959; Lorenzo et al., 2014). Antelope jackrabbits appear to tolerate

nonnative grass species, such as Lehmann lovegrass (Eragrostis lehmanniana), while

other jackrabbits, such as the Tehuantepec jackrabbit, prefer native vegetation (Farías and

Fuller, 2009). Other small mammal communities change in response to Lehmann

lovegrass, depending on their preference for dense vegetation (Litt and Steidl, 2011).

Additionally, antelope jackrabbits do not need free water to survive, but will drink water

if offered (Vorhies and Taylor, 1933). Groups of up to 8 individuals have been

documented around existing cattle tanks (pers. obs.), but the importance of available

water is not known for this species.

Buenos Aires National Wildlife Refuge (BANWR) in southcentral Arizona

provides a unique study area where the landscape is greatly altered due to cattle grazing,

55

leading to shrub encroachment and increase in non-native vegetation. Although the land

has been anthropogenically-altered, cattle are no longer permitted on the refuge and

management goals include the restoration of the semidesert grassland to a pre-settlement

state with the use of prescribed burns, native seed planting, and mechanized removal of

trees (USFWS, 2012). Our goals of this study were to 1) better understand antelope

jackrabbit habitat characteristics, including structure and composition, in a modified

landscape, and 2) determine if a spatial relationship exists between the antelope

jackrabbit and the Pima pineapple cactus.

STUDY AREA

This study was conducted on the Buenos Aires National Wildlife Refuge

(31.5501° N, 111.5507° W), in Pima County in southcentral Arizona, 90 km southwest of

Tucson, Arizona. BANWR encompasses 46,450 ha in the Altar Valley basin. The U.S.-

Mexico international border is the southern boundary of BANWR, with the Baboquivari

Mountains bordering on the west and the Sierrita Mountains bordering on the east.

Elevation ranges between 925 to 1,400 m (Koenen and Krausman, 2002), however most

of the refuge occurs within 950 to 1,150 m (McLaughlin, 1992). BANWR is mainly

comprised of semidesert grassland and Sonoran desertscrub, with some historical hints of

a subtropical Sonoran savanna grassland (Brown 1994). Exotic grasses (Eragrostis

chloromelas, E. lehmanniana, Sorghum halepense), shrubs (Ambrosia dumosa, Baccharis

sarothroides, Gutierrezia sarothrae), native grasses (Aristida hamulosa, A. ternipes,

Bouteloua rothrocku, B. repens, Digitaria californica), cactus species (Carnegiea

gigantea, Echinocereus fasciculatus, Opuntia spp.) and velvet mesquite stands can be

found on the refuge (McLaughlin, 1992; Brown, 1994), however, much of the semidesert

56

grassland is now dominated by Lehmann lovegrass (McLaughlin, 1992). BANWR

receives an average of 410 mm annual precipitation (Sellers et al., 1985), with monsoon

rains occurring in July and August and winter rains occurring between December and

March (Arizona-Sonora Desert Museum, 2000; McLaughlin, 1992).

METHODS

Home Range Size

We used hoop nets and spot lights to capture animals at night. When we caught

an individual, we collected biological data including weight, sex, and reproductive status

and fit adult animals with a VHF radio collar (Wildlife Materials, Murphysboro, IL;

Altemus 2016). We used single-person biangulation to locate animals at least once per

week. We used Backpacker GPS Trails (Trimble Navigation Ltd., Sunnyvale, CA) to

record locations. We used the “best biangulation” function in Location of a Signal

(LOAS; Ecological Software Solutions LLC, Hegymagas, Hungary) to estimate animal

locations. We used the Spatial Analyst (Environmental Systems Research Institute Inc.

[ESRI], Redlands, CA) and Animal Movement extensions (Hooge and Eichenlaub, 1997)

in ArcView 3.3 to calculate 95% (home range) fixed kernel density estimates (Laver and

Kelly, 2008). All field work was conducted in accordance with the American Society of

Mammalogists guidelines and approved by the University of Arizona’s Institutional

Animal Care and Use Committee (Sikes et al., 2011; IACUC protocol #13-480). Field

methods were conducted under permits from the Arizona Game and Fish Department

(LIC #SP654189) and Buenos Aires National Wildlife Refuge (Special Use Permit

#2013-028).

Vegetation Structure and Composition

57

Data collection

We collected data on vegetation structure and composition at vegetation points

within antelope jackrabbit home ranges and at random points within the study area to

determine habitat selection for home ranges at the population level (type II study design:

Manly et al., 2002; second order resource selection: Johnson, 1980). We collected

vegetation measurements at no fewer than 10 points within each animal’s home range;

these represented use. We also collected vegetation measurements at randomly generated

points in space available to antelope jackrabbits; these points represented availability.

We estimated habitat available to antelope jackrabbits by creating one minimum convex

hull polygon in the study area around all antelope jackrabbit telemetry fixes using the

Rectangle tool in ArcMap. Within the polygons, we generated 225 random vegetation

points (“available points”). We located vegetation points in the field using an eTrex 10

GPS device (Garmin International, Inc., Olathe, KS).

We used the visual obstruction method to measure vegetation structure (Robel et

al., 1970). We constructed a 1.2 m Robel pole with 2.5 cm bands up to 60 cm. We

attached a 4 m-long string at 1 m to standardize the measurement height and distance.

We recorded the lowest band that was visible from the ground from each of the four

cardinal directions to determine the visual obstruction measurement (VOM) at each

vegetation point. For data interpretation purposes, VOMs are converted to cm by

multiplying VOM by 2.5.

We recorded the dominant vegetation characteristic within a 20 m radius

surrounding the vegetation point to measure vegetation composition. We observed 11

vegetation characteristics: bare ground, road, wash, kangaroo rat mound, Lehmann

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lovegrass, Johnson grass, grass species (not Lehmann lovegrass or Johnson grass), desert

broom, mesquite (Prosopis spp.), acacia (Acacia spp.), palo verde (Parkinsonia spp.),

snakeweed, forbs, saltbush (Atriplex spp.), wolfberry (Lycium spp.), or cactus (Family

Cactaceae).

Data analysis

We averaged the four cardinal direction VOM readings to get mean VOM for

each vegetation point. We averaged VOMs at all vegetation points in each animal’s

home range to get the mean VOM for each animal. We compared VOMs between use

and available points with t-tests. We calculated a simple linear regression to predict

home range size based on VOM.

To determine if a difference existed between vegetation characteristics at use

points and available points, we conducted Pearson’s Chi-square tests. First, we pooled

the original 11 vegetation characteristics into 6 vegetation groups: forbs, bare ground,

woody plants, nonnative grasses (Johnson grass and Lehmann lovegrass), native grass,

and trees. We determined the dominant vegetation characteristic for each vegetation site.

We summed the number of vegetation sites with each dominant characteristic; this was

our observed value for that dominant characteristic. We did the same with available sites

to determine the expected values for each dominant characteristic.

Pima Pineapple Cactus Locations

We assessed proximity of antelope jackrabbit locations to PPC locations. Pima

pineapple locations were recorded by GPS by U.S. National Park Service field

technicians during stratified random sampling of vegetation plots across BANWR in the

summers of 2012 and 2013. Antelope jackrabbit telemetry (use) points collected between

59

December 2013–November 2015 were compared to random (available) points within the

study area. We compared the average distance of telemetry points to PPC and average

distance of random points to PPC using a Welch’s t-test. We used the Near tool in

ArcMap to determine which points were within 193.5 m (the radius of our smallest

antelope jackrabbit home range) of a PPC; these points were considered “near” a PPC.

We conducted a Fisher’s exact test to compare the number of telemetry points near PPC

to number of available points near PPC. We used a t-test to compare telemetry point-

PPC distances to available point-PPC distances.

We separated antelope jackrabbit telemetry points temporally and created a subset

of locations during the PPC fruiting season from August to December. We used a t-test

to compare mean distance of antelope jackrabbit locations to PPC during the fruiting

season to mean distance of available points to the nearest PPC.

Water Tanks

We determined proximity of antelope jackrabbit locations to water tanks.

BANWR staff provided us with locations of some water tanks; we located other water

tanks with the help of an Arizona Game and Fish Department map (AZGFD and Klima

2012) and with Google Maps. We used the Near tool in ArcMap to measure distances of

each telemetry point to the nearest water tank and distances of random points to the

nearest water tank. We used a t-test to compare the average distance of antelope

jackrabbit telemetry points to water tanks and the average distance of random points to

water tanks.

RESULTS

Home Range Size

60

Mean ± SD number of telemetry locations for home range estimates was 52.00 ±

23.26. Mean ± SD home range size was 30.07 ± 16.85 ha.

Vegetation Structure and Composition

We measured vegetation structure and composition at 104 use points in 10

antelope jackrabbit home ranges (mean 10.40 ± 0.52 vegetation points per home range).

We measured an average of 0.43 ± 0.22 vegetation points per home range hectare.

Measurements were taken from October–December 2015.

Mean VOM for use points in individual antelope jackrabbits home ranges ranged

between 3.75–27.22 cm, with a mean of 12.68 ± 8.87 cm (Figure 1). Mean VOM for

available points was 13.04 ± 13.57 (Figure 1). VOMs at all use points and all available

points did not differ (t240 = 0.10, P = 0.92). Home range size and VOM were not related

(F1, 8 = 0.81, P = 0.39, R2 = 0.09).

Vegetation groups between all use points and all available points did not differ

(Table 1; χ52 = 7.90, P = 0.16). Individual vegetation characteristics did not differ (P =

1.0; Figure 2). Antelope jackrabbits tended to use slightly more Lehmann lovegrass than

available and slightly less mesquite than available (Figure 2).

Pima Pineapple Cactus Locations

Telemetry points (n = 730) of 22 radio-collared antelope jackrabbits represented

use points and random locations (n = 225) within the study area represented available

points. We had 115 PPC locations. The average distance of random points to the closest

PPC (919.13 ±718.72 m) was smaller than the distance of telemetry points to the closest

PPC (1186.25 ± 846.93 m) (t432 = -4.67, P < 0.01). Of telemetry points, 11.10% (n = 81)

were near (within 193.5 m) a PPC and of random locations, 11.11% (n = 25) were near a

61

PPC. The percent of telemetry points and random locations near a PPC did not differ (P

= 1.0). Of near telemetry points and near random points, the mean ± SD distance of

telemetry points to a PPC was 121.20 ± 51.20 m and for random points near a PPC, the

mean distance was 120.70 ± 49.71 m. Distances of antelope jackrabbit locations near

PPC and distances of random locations near PPC did not differ (t104 = 0.04, P = 0.97).

Mean distance of antelope jackrabbit locations to PPC during the fruiting season was

984.52 ± 795.76 m; this did not differ from random locations to PPC (t423 = 0.89, P =

0.37).

Water Tanks

Mean distance of antelope jackrabbit telemetry locations (n = 730) to water tanks

(n = 22) was 1153.40 ± 656.84 m and mean distance of random points (n = 225) to water

tanks was 1206.37 ± 571.48 m. Mean distance of antelope jackrabbit telemetry locations

and random points to water tanks did not differ (t953 = -1.09, P = 0.28).

DISCUSSION

Antelope jackrabbits selected similar vegetation structure and composition to

what was available. We did not find a spatial association between antelope jackrabbit

locations and PPC throughout the year, or during the PPC fruiting season.

Vegetation Structure

Jackrabbits require heterogeneity in vegetation height and species diversity to

provide food resources and allow for the detection and evasion of predators (Lechleitner,

1958; Longland, 1991; Marín et al., 2003; Smith et al., 2004). High VOMs are positively

correlated with vegetation biomass and vegetation density (Robel et al., 1970; Uresek and

Benzon, 2007). VOMs at use points were similar to VOMS at available points,

62

indicating that antelope jackrabbits inhabit areas that are similar to the surrounding and

available area. Home range size could not be predicted with VOM, which indicates that

factors besides resource quality and availability may influence home range size, to

include population dynamics or predation. Intraspecific and interspecific competition can

influence habitat structure indirectly by altering available biomass (Pulliainen and

Tunkkari, 1987). Black-tailed jackrabbits and desert cottontails (Sylvilagus audubonii)

are sympatric with antelope jackrabbits on BANWR, however interactions and

abundance and density of these species is not known. Vegetation structure is also linked

to predation risk for hares (Jennings et al., 2003; Smith et al., 2004, 2005). Although

predator abundance and diversity is unknown on BANWR, active bobcat and coyote

hunting occurs on various ranches surrounding BANWR. If predators are scarce, prey

species like the antelope jackrabbit, to require less vegetative cover. Determining

predator dynamics on BANWR would help determine contributing variables that

influence the differences in habitat selection by antelope jackrabbits.

Vegetation Composition

Vegetation characteristics were similar at used and available vegetation points.

Antelope jackrabbits tended to use slightly more Lehmann lovegrass than available.

Despite negative connotations associated with non-native species, for some species, bird

abundance and density in areas with native and non-native grasses does not differ,

indicating that some species select habitat based on structure rather than composition

(e.g., Lloyd and Martin, 2005; Flanders et al., 2006; Kennedy et al., 2009; Thompson et

al., 2009). Another desert grassland jackrabbit species, the white-sided jackrabbit (Lepus

callotis), is negatively correlated with woody shrubs, illustrating that structure is an

63

important habitat determinant (Nelson, 1909; Dunn et al., 1982; Bednarz and Cook, 1984;

Findley, 1987; NMDGF and Traphagen, 2011). Antelope jackrabbits also tended to use

slightly more mesquite than available, which is in accordance with other findings that

specify velvet mesquites are the dominant overstory plant where antelope jackrabbits are

found (Brown et al., 2014). Our results indicate that antelope jackrabbits are capable of

inhabiting a variety of environments with diverse plant species, placing importance on

vegetation structure.

Pima Pineapple Cactus Spatial Association

Herbivores are important seed dispersers (Janzen, 1984) and rabbit pellets often

contain germinable seeds (Pijl, 1972; Welch, 1985; D’Antonio, 1990; Zedler and Black,

1992). Lagomorphs are seed dispersal agents for cactus species (Cook, 1942; Timmons,

1942), including the PPC (WestLand Resources, Inc., 2009, unpublished report). We did

not find evidence of a spatial relationship between PPC and antelope jackrabbits,

however, our study observed a subset of antelope jackrabbits and PPCs and may not be

representative of the entire population. Antelope jackrabbits may eat PPC fruit

opportunistically rather than seek this uncommon and seasonal food out. Telemetry

locations during the fruiting season, from August to December (Roller, 1996), were not

closer to PPC than random points to PCC. Further studies observing more fine-scale

movements may help to clarify the relationship between antelope jackrabbits and PPC.

Importance of Water

In an arid landscape with uncertain and irregular precipitation events, water

availability can be beneficial to animals (Rosenstock, 1999). Some animals, like the

antelope jackrabbit, do not need free water to survive, but will drink water if it is

64

available (Vorhies and Taylor, 1933). Antelope jackrabbits did not have an affinity for

water tanks, however animals may congregate around tanks for mating purposes or

seasonally as water is more scarce (Elder, 1956). Future research focusing on

movements around water sources would help identify what role these landscape features

play for antelope jackrabbits.

An Altered Landscape

BANWR, a former rangeland, has transformed into a nonnative grass-dominated

shrubland. In other areas of former antelope jackrabbit range, habitat has been converted

to other land uses; farmland and urbanization has taken over about 20% of the antelope

jackrabbit’s range (Brown et al., 2014). Aside from local landscape change, regional and

global patterns have the potential to impact wildlife as well. The southwest region

already shows warming trends throughout the Sonoran Desert (Weiss and Overpeck,

2005) and is expected to experience increased warming and drought, leading to increased

wildfires (U.S. Global Change Research Program, 2014). The region is also expected to

experience a declining water supply, and increased flooding and erosion (U.S. Global

Change Research Program, 2014), all of which may impact antelope jackrabbits and the

Pima pineapple cactus.

IMPLICATIONS

A specified antelope jackrabbit hunting season or bag limit does not exist in

Arizona, the only U.S. state where antelope jackrabbits occur. Furthermore, if antelope

jackrabbits are necessary for the dispersal or propagation of the PPC, the need for

research on antelope jackrabbit’s ecological relationships with other lagomorphs,

65

resources, and threatened species may increase and will help inform land management

decisions regarding grazing, prescribed burns, and native seed planting.

ACKNOWLEDGMENTS

Thank you to the Buenos Aires National Wildlife Refuge administrative and

biological staff and National Park Service for allowing us access to the PPC data and

thanks to the numerous technicians who gathered this data. We are appreciative of K.

Bennett, J. Derbridge, and E. Goldstein for revisions and comments. Special thanks to

M. Merrick for providing feedback and assisting with GIS analysis. Thank you to A.

Moreno for his assistance with vegetation measurements.

This work was supported by the Reid Park Zoological Society, T&E Inc.,

USFWS, National Park Service (NPS), and Desert Southwest Cooperative Ecosystem

Studies Unit (DSCESU), Arizona Agricultural Experiment Station, University of Arizona

Vice President for Research, Mt. Graham Red Squirrel Research Program, Koprowski

Conservation Research Laboratory, and the American Society of Mammalogists.

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Figure 1. Mean ± SE vegetation obstruction measurements (VOMs) in cm at antelope

jackrabbit (Lepus alleni) use and available vegetation points on Buenos Aires National

Wildlife Refuge, Arizona taken between October–December 2015.

78

Figure 2. Differences between the average percentages of each dominant vegetation

characteristic at used vegetation points and available vegetation points on Buenos Aires

National Wildlife Refuge, Arizona for antelope jackrabbit (Lepus alleni). Abbreviations

are amaranth (AMAR), bare ground (BARE), cactus (CACT), desert broom (DEBR),

forbs (FORB), native grass species (GRAS), Johnson grass (JOHN), Lehmann lovegrass

(LEHM), mesquite species (MESQ), palo verde (PALV), snakeweed (SKWD), and

wolfberry (WOLF). Measurements were taken between taken between October–

December 2015.

79

Table 1. Mean percentage of each dominant vegetation group at use points in antelope

jackrabbit (Lepus alleni) habitat and at available points for all locations on Buenos Aires

National Wildlife Refuge, Arizona, measured October–December 2015.

Forb

group

Bare

ground

Woody

group

Grass

group

Nonnative

grass group

Tree

group Total

All Use 7.7 1.9 1.9 19.2 56.7 12.5 99.9

Available 5.0 6.5 4.3 19.6 49.3 15.2 99.9

80

APPENDIX C: Supplemental data on antelope jackrabbit (Lepus alleni) captures, core

areas, home ranges, and seasonal ranges

The following tables contain capture and radio collar data, raw data of core area

estimates (50% fixed kernel), home range estimates (95% fixed kernel), seasonal range

estimates (ha), and seasonal home range usage percentages for antelope jackrabbits

(Lepus alleni) on Buenos Aires National Wildlife Refuge, Arizona between December

2013–November 2015.

81

Table 1. Capture history for all antelope jackrabbits caught on Buenos Aires National

Wildlife Refuge, Arizona, between December 2013–November 2015. Fates are of

collared animals and indicate whether the collars were found (mortality), whether the

collars stopped transmitting a signal (missing), or whether the animal’s collar was still

transmitting a signal at the end of the study (alive).

An

ID

Date

Caught

Release Status Sex Repro

Status

Age Weight

(kg)

Fate as of

5/9/16

1 02-Nov-13 Capture

mortality

Female Non-repro Sub-

adult

2.25

2 08-Nov-13 Escaped,

without collar

Unknown Unknown Adult

3 08-Nov-13 Escaped,

without collar

Unknown Unknown Adult

4 16-Nov-13 OK, with

collar

Unknown Unknown Adult Mortality

5 07-Dec-13 OK, with

collar

Female Non-repro Sub-

adult

Alive

6 28-Dec-13 OK, with

collar

Male Non-repro Adult 3.42 Missing

7 30-Dec-13 OK, with

collar

Unknown Unknown Adult 3.91 Mortality

8 16-Feb-14 OK, without

collar

Unknown Non-repro Leveret 0.96

9 25-Feb-14 OK, without

collar

Unknown Non-repro Leveret 0.25

10 25-Mar-14 OK, without

collar

Male Partially

scrotal

Sub-

adult

1.24

11 28-Mar-14 OK, without

collar

Female Non-repro Sub-

adult

1.48

12 03-Apr-14 OK, without

collar

Female Non-repro Sub-

adult

1.16

13 04-Apr-14 OK, without

collar

Unknown Non-repro Leveret 0.24

14 04-Apr-14 Other Unknown Non-repro Leveret 0.21

15 18-Apr-14 OK, with

collar

Male Scrotal Adult 3.77 Mortality

16 18-Apr-14 OK, with

collar

Male Scrotal Adult 3.74 Mortality

17 18-Apr-14 OK, without

collar

Male Partially

scrotal

Leveret 0.71

82

18 10-May-14 OK, without

collar

Female Non-repro Sub-

adult

1.51

19 10-May-14 OK, with

collar

Female Non-repro Sub-

adult

1.91 Missing

20 13-Jun-14 Capture

mortality

Male Partially

scrotal

Adult

21 18-Jun-14 OK, with

collar

Male Non-repro Adult 5.01 Alive

22 25-Jun-14 Escaped,

without collar

Unknown Unknown Adult

23 26-Jun-14 Escaped,

without collar

Unknown Unknown Sub-

adult

24 02-Jul-14 OK, with

collar

Female Nipples

visible

Adult 4.29 Mortality

25 03-Jul-14 OK, without

collar

Female Non-repro Sub-

adult

1.61

26 15-Jul-14 OK, without

collar

Female Non-repro Sub-

adult

1.45

27 22-Jul-14 OK, with

collar

Male Scrotal Adult 2.85 Mortality

28 23-Jul-14 OK, with

collar

Male Partially

scrotal

Adult 2.87 Mortality

29 04-Sep-14 OK, with

collar

Female Non-repro Sub-

adult

1.94 Alive

30 19-Sep-14 OK, with

collar

Female Non-repro Sub-

adult

1.65 Mortality

31 19-Sep-14 OK, without

collar

Female Non-repro Sub-

adult

1.24

32 25-Sep-14 OK, with

collar

Female Nipples

visible

Adult 3.61 Mortality

33 25-Sep-14 OK, with

collar

Female Non-repro Adult 2.90 Missing

34 27-Sep-14 OK, with

collar

Female Non-repro Adult 3.16 Missing

35 09-Oct-14 OK, with

collar

Male Scrotal Adult 3.71 Mortality

36 21-Oct-14 OK, with

collar

Female Recent

lactating

Adult 3.91 Mortality

37 22-Nov-14 OK, with

collar

Female Non-repro Adult 2.99 Alive

38 12-Feb-15 OK, with

collar

Female Non-repro Adult 3.49 Mortality

39 12-Feb-15 Other Female Non-repro Leveret 0.52

40 20-Mar-15 OK, with

collar

Female Lactating Adult 3.92 Mortality

83

41 03-Jun-15 OK, with

collar

Male Scrotal Adult Alive

Table 2. Number of antelope jackrabbits (Lepus alleni) caught and collared, by sex, on

Buenos Aires National Wildlife Refuge, Arizona, between December 2013–November

2015.

Male Female Unknown Total

Captured 11 20 10 41

Radio-collared 8 12 2 22

Table 3. Number of antelope jackrabbits (Lepus alleni) caught and collared, by age

group, on Buenos Aires National Wildlife Refuge, Arizona, between December 2013–

November 2015.

Adult Subadult Leveret Total

Captured 22 13 6 41

Radio-collared 18 4 0 22

84

Table 4. Core area (50% fixed kernel), home range (95% fixed kernel) and seasonal range

(95% fixed kernel) density estimates (ha) for individual antelope jackrabbits (Lepus

alleni) between December 2013–November 2015 on Buenos Aires National Wildlife

Refuge, Arizona. Winter months were December, January, February and March. Spring

months were April, May, and June. Monsoon months were July, August, and September.

Fall months were October and November.

Animal

ID

Study

area

Core

area

Home

range

Seasonal:

winter

Seasonal:

spring

Seasonal:

monsoon

Seasonal:

fall

F5 North 1.36 14.54 14.36 11.90 16.18 1.89

F24 South 1.94 17.46

F29 South 3.74 24.72 7.11 9.73 39.99 9.78

F37 North 4.19 22.24 17.24 13.61 12.63 27.41

F40 North 3.95 18.91

Female

mean

3.04 ±

1.29

19.57 ±

3.99

12.90 ±

5.22

11.75 ±

1.94

22.93 ±

14.88

13.03 ±

13.07

M6 North 5.96 38.36 36.17 42.00 38.24 25.29

M15 North 4.68 43.57 41.65 40.69 39.31 29.68

M16 North 4.22 24.02 21.52 25.56 14.41 4.54

M21 South 2.04 11.76 5.41 9.78 11.08 9.39

M27 South 14.68 67.48 16.03 78.45 55.59 14.50

M35 North 10.96 36.57

M41 South 4.70 30.04

Male

mean

6.75 ±

4.44

35.97 ±

17.42

24.16 ±

14.79

39.30 ±

25.51

31.73 ±

18.68

16.68 ±

10.59

Overall

mean

5.20 ±

3.87

29.14 ±

15.58

19.94 ±

12.91

29.09 ±

23.98

28.47 ±

16.90

15.44 ±

10.78

85

Table 5. Home range usage percentage for each season and degree of interseason

similarity (DIS) for all seasons for each antelope jackrabbit (Lepus alleni) from

December 2013–November 2015 on Buenos Aires National Wildlife Refuge, Arizona.

Home range usage percentage is the seasonal home range size is divided by the home

range. Lower DIS values indicate a more sedentary lifestyle, whereas higher DIS values

are indicative of actively moving individuals. Winter months were December, January,

February and March. Spring months were April, May, and June. Monsoon months were

July, August, and September. Fall months were October and November.

Animal

ID Winter Spring Monsoon Fall DIS

F5 69.82 % 70.33 % 88.28 % 13.03 % 14.97 %

M6 76.76 % 91.87 % 82.71 % 52.47 % 48.72 %

M15 54.81 % 78.48 % 71.54 % 51.95 % 34.97 %

M16 73.27 % 84.13 % 57.01 % 18.89 % 24.65 %

M21 42.28 % 76.77 % 84.31 % 57.18 % 28.81 %

M27 19.51 % 71.39 % 59.43 % 18.87 % 19.02 %

F29 28.74 % 39.36 % 80.40 % 39.30 % 15.10 %

F37 62.31 % 56.57 % 52.55 % 76.48 % 32.07 %

Mean 53.44 ±

21.29 %

71.11 ±

16.50 %

72.03 ±

13.96 %

41.02 ±

22.48 %

27.29 ±

11.46 %