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The UPDATE published this issue in partnership with the Defenders of Wildlife. Defenders of Wildlife is a leading nonprofit con- Endangered Species servation organization recognized as one of the nation's most progressive UPDATE advocates for wildlife and its habitat. Defenders uses education, litigation, A forum for information exchange on research and promotion of sound conservation policies to protect wild ani- endangered species issues JulyIAugust 2001 Vol. 18 No. 4 mals and plants in their natural communities. Defenders has been a national leader in restoration and protection for two decades. Founded in 1947, M. Elsbeth McPhee ...... Managing Editor Jennifer Jacobus Mackay...... Associate Editor defenders is a 501(c)(3) membership organization with more than 430,000 Nina Fascione...... Guest Editor members and supporters. It is headquartered in Washington, DC with field Stephen Kendrot ...... Guest Editor offices around the country. Martin E. Smith...... Guest Editor Paula Zemrowski ...... Content Editor Last November, Defenders of Wildlife hosted Carnivores 2000, a con- Deborah Kleinman...... Content Editor ference on carnivore conservation in the twenty-first century. The confer- Chris Wood...... Content Editor ence, held in Denver, , spanned the full taxonomic range of carni- SaulAlarcon...... Web Page Coordinator Terry L. Root ...... Faculty Advisor vores and addressed both biological and sociological issues impacting car- nivore conservation in and abroad. Due to its overwhelming Advisorv Board Richard Block success, Defenders of Wildlife will hold a second conference, Carnivores Santa Barbara Zoological Gardens 2002, in Monterey, California November 18 to 20,2002. This conference will Susan Haig likewise encompass the full range of marine and terrestrial carnivores. For more Forest and Rangeland Ecosystem Science Center, USGS information, visit www.defenders.org/carnivores2002. We hope to see you there. Oregon State University Chris Howes Chicago Zoological Society Patrick O'Brien Chevron Ecological Services Hal Salwasser U.S. Forest Service, Boone and Crockett Club

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Cover: Black ( americanus). Photograph by B. Moose Peterson~WRP.

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94 Endangered Species UPDATE Vol. 18 No. 4 2001 Table of Contents

Carnivore Conservation in the Twenty-first Century...... 97 Nina Fascione and M. Elsbeth McPhee

I. Theory and Methods in Carnivore Conservation Who Limits Whom: Predators or Prey? ...... 98 Roger A. Powell

A New Approach to Understanding Canid Populations Using an Individual-Based Computer Model: Preliminary Results...... 103 William C. Pitt, Frederick F. Knowlton, and Paul W. Box

11. Carnivores and Habitat Mapping Occupied Habitat for Forest Carnivores in the American West and Estimating their ...... 107 David L. Gaillard

Road Density as a Factor in Habitat Selection by and Other Carnivores in the Great Lakes Region...... 110 Adrian P. Wydeven, David J. Mladenoff, Theodore A. Sickley, Bruce E. Kohn, R. P. Thiel, and Jennifer L. Hansen

A Predator-Habitat Assessment for Felids in the Inland Atlantic Forest of Eastern Paraguay: A Preliminary Analysis ...... 115 Gerald L. Zuercher, Philip S. Gipson, and Kim Hill

111. Raptor Conservation Status of the California Condor and Mortality Factors Affecting Recovery...... 120 Kelly J. Sorenson, L. Joseph Burnett, and James R. Davis

Raptors as Vermin: A History of Human Attitudes towards Pennsylvan~aI s B~rds. of Prey...... 124 Keith L. Bildstein

IV. Felid Conservation Status and Conservation of Endangered along the US.-Mexico Border...... 129 Melissa Grigione, Arturo Caso, Rurik List, and Carlos Lopez-Gonzalez

Molecular Scatology as a Conservation Tool ...... 133 Laura Farrell

Linking Snow Leopard Conservation and People-Wildlife Conflict Resolution: Grassroots Measures to Protect the Endangered Snow Leopard from Herder Retribution...... 138 Rodney Jackson and Rinchen Wangehuk

Vol, 18 No. 4 2001 Endangered Species UPDATE 95 K Canid Conservation The Status of the Wolf Population in Post-Soviet Kyrgyzstan...... 142 CJ Hazell

Gray wolf restoration in the Northwestern United States ...... 147 Ed Bangs, Joe Fontain, Mike Jimenez, Tom Meier, Carter Niemeyer, Doug Smith, Kerry Murphy, Deb Guernsey, Larry Handegard, Mark Collinge, Rod Krischke, John Shivik, Curt Mack, lssac Babcock, Val Asher, and Dominic Domenici

The Feasibility of Gray Wolf Reintroductionto the Grand Canyon Ecoregion...... 153 Paul G. Sneed

Canis Soupus: Eastern Wolf Genetics and its Implications for Wolf Recovery in the Northeast United States ...... 159 Nina Fascione, Lisa G. L. Osborn, Stephen R. Kendrot, and Paul C. Paquet

Captivity, Inbreeding, Cross-Lineage Matings, and Body Size in Mexican Wolves ...... 164 Richard Fredrickson and Philip Hedrick

First Swift , velox, Reintroduction in the USA: Results of the First Two Years ...... ,167 Clio Smeeton and Ken Weagle

lnterspecific Interactions Among Wild Canids: Implications for the Conservation of Endangered San Joaquin Kit ...... 171 Brian L. Cypher, Howard 0. Clark, Jr., Patrick A. Kelly, Christine Van Horn Job, Gregory D. Warrick, and Daniel F. Williams

VI. Predation Management Chemical Repellents and Other Aversive Strategies in Predation Management...... 175 J.R. Mason, J.A. Shivik, and M.W. Fall

Effectiveness of Livestock Guarding for Reducing Predation on Livestock...... I82 William F. Andelt

VII. Opinions Historic Attitudes and Images and the Implications on Carnivore Survival...... 186 Karlyn Atkinson Berg

Bringing Down the Walls ...... 190 Gale H. Ford

News from Zoos...... 1 9 1

96 Endangered Species UPDATE Vol. 18 No. 4 2001 Carnivore Conservation in the Twenty-first Century Nina Fascione Defenders of Wildlife, 1101 14th St. NW, Suite 1400, Washington DC 20005; [email protected] M. Elsbeth McPhee Endangered Species UPDATE, University of Michigan, 430 E. University, Ann Arbor, MI 481 09; (734) 763-3243; (fax) (734) 936-2195; [email protected]

The dawn of this new millennium is Mexico-U.S. border. Smeeton and conservation. Finally, the rigorous proving to be an interesting time for Weagle (p. 167) describe successful application of ecological theory to carnivores. In some areas predators reintroductions of the swift fox conservation questions has lead to - seem to be rebounding after years of (Vulpes velox) into the of greater understanding of population declining populations resulting from North America. dynamics and behavior within imper- habitat loss and human persecution. In addition, new technologies iled carnivore populations (see For example, in this volume, Bangs have fostered improved research in Powell p. 98; Zuercher p. 115; et al. (p. 147) discuss the remarkable both field studies and laboratory set- Fredrickson and Hedrick p. 164; and recovery gray wolves ( lupus) tings. Improvements in radio-telem- Cypher et al. p. 171). Understanding are making in the lower 48 states, and etry, remote-sensing work, geo- issues such as genetics, natural his- Sneed (p. 153) describes plans to re- graphic information systems (see tory, habitat needs, and predator-prey store wolves into the Grand Canyon Gaillard p. 107 and Wydeven et al. p. relationships will enable scientists to ecoregion. Grigione et al. (p. 129) 1lo), genetics (see Farrell p. 133 and manage and conserve carnivore popu- report signs of jaguars ( Fascione et al. p. 159), and computer lations well into the future. onca), ocelots ( pardalis), modeling (see Pitt et al. p. 103) have Unfortunately, not all carnivores and (Herpailurus led to increased knowledge of the are showing such positive signs of yaguarondi) on both sides of the myriad factors affecting carnivore recovery (see Sorenson p. 120 and Hazel1 p. 142), and all predators still face innumerable threats. Habitat Subscribe to the UPDATE today! loss, competition with humans for resources and human persecution are In its 18 years of publication, the Endangered Species UPDATE, published by some of the major issues with which the School of Natural Resources and Environment at the University of Michi- wildlife managers must contend. gan, has established itself as the primary forum for government agencies, Anti-predator sentiment remains conservation organizations, private consulting and law firms, zoos, museums, educational institutions, and others to exchange ideas and information on strong in some arenas as well (see endangered species issues. Bildstein p. 124; Jackson p. 138; Berg p. 186; Ford p. 190;Mason et al. 175; -Subscriptionrates are: regular subscriptions $28 and Andelt p. 182). studentlsenior (65+) $23 address outside the US add $5 Continuing to expand our knowl- electronic subscription $20 edge of carnivores will be essential to combating these issues and find- To subscribe, make check payable to the Universityof Michigan and mail to: ing new and innovative ways to en- Endangered Species UPDATE able humans to co-exist with healthy School of Natural Resources and Environment carnivore populations. Forums such University of Michigan 430 E. University as the Defenders of Willdife's confer- Ann Arbor, MI 481 09-1115 ences (Carnivores 2000 Denver, CO November 2000; and Carnivores or visit our website: http://www.umich.edu/-esupdatel 2002 Monterey, CA November 2002), and the Endangered Species UPDATE are vital to the continued success of carnivore conservation ef- forts. We thank the authors for their contributions to this edition and their work on carnivore conservation.

Vol. 18 No. 4 2001 Endangered Species UPDATE 97 Theory and Methods in Carnivore Conservation Who Limits Whom: Predators or Prey?

Roger A. Powell Department of Zoology, North Carolina State University, Raleigh, North Carolina 27695-7617

Abstract populations can be limited by the availability of food (limited from the bottom of the food chain up), by predators (limitedfrom the top of the food chain down), or by the interaction of these two processes. Whether carnivores, in particulal; limit the populations of their prey, or are limited by those prey, has long been controversial and is critical to conservation of prey and predatory species. I return to the question because it is a good question that we wish to have answered and, in part, because it has no simple answer. Our knowledge of ecological communi- ties has matured to the point that we can tease the question apart, look at its pieces, andfind conditional answers. Predator-prey models suggest that predators may limit prey populations on one scale while food limits prey, and prey limit predators on another scale. Predator populations are always limited by the availability of their prey. Data from the literature supports action on two time scales.

Introduction Strong 1996; Strong 1992; White Here, I ignore the abilities of plants "Who limits whom: predators or prey?" 1978). From this perspective, preda- to defend themselves and explore the is an old question that resurfaces regu- tors in species-rich, terrestrial commu- effects on herbivores and predators of larly because, in part, it is a good ques- nities rarely have large effects on the varying the productivity of plants that tion that we wish to have answered and, populations of their prey but, instead, is available to herbivores. For conve- in part, because it has no simple an- are limited by them. Herbivore popu- nience, I call this available productiv- swer. I address the question once again lations limit the populations of their ity of plants "primary productivity." because ow knowledge of ecological predators because predator populations communities has matured to the point are small and herbivores are well Model predators and prey that we can tease the question apart, adapted to avoid their predators. Using a difference equation model look at its pieces, and find conditional In contrast, a Top-Down perspec- similar to that used by Boyce and answers. These answers provide a ba- tive was outlined by Elton (1927), Anderson (1999), incorporating Type sis for managing populations of preda- then made more rigorous by Hairston 3 functional responses (Holling 1959) tors and prey, for conserving endan- et al. (1960) in their "The World is and density-dependent population gered species, for understanding what Green" paper, and has been explored growth for both predators and prey, I kinds of population fluctuations we can extensively by Fretwell, Oksanen and developed a system with three trophic affect and what kinds we can not, and coworkers (Fretwell 1977 ,1987; levels. The responses of this system to for knowing where in an ecological sys- Oksanen et al. 1981; Oksanen and changes in primary productivity re- tem our management energies should Oksanen 2000) as the Exploitation semble the classic Top-Down trophic be targeted. Ecosystem Hypothesis. From this pattern (Figure 1a) except that the her- The debate regarding who limits perspective, plants are vulnerable to bivore graph has a "hump" (Figure lb). whom has been viewed from two pre- folivores but are seldom severely de- The model predicts that at high levels dominant perspectives. The Bottom- foliated because of predation pressure of primary productivity, predators limit Up perspective emphasizes that plant on herbivores. The Exploitation their prey to lower population sizes than defenses against herbivores and herbi- Ecosystem Hypothesis is not lim- they do at intermediate levels of pri- vore defenses against predators are so ited to productive, species-rich sys- mary productivity. Similarly, when her- strong that communities are limited at tems and is most often tested using bivores are not limited by predators, each level predominantly by food communities with vertebrate preda- they limit the standing crop of vegeta- (Mwdoch 1966; Polis 1999; Polis & tors and prey. tion. The model also predicts a most

98 Endangered Species UPDATE Vol. 18 No. 4 2001 basic characteristic of herbivore and predators to exist. If primary produc- wolves and Man (Gray 1987; Mech predator populations: primary produc- tivity is not great enough, herbivore 1992). In the Upper Midwest states, tivity establishes the conditions that populations are not large enough to wolves and white-tailed allow herbivores, or herbivores and support predators and predator popu- (Odocoileus virginianus) form the lations can not be- dominant predator-prey system come established. (Mech 2000). TOPeDOWN TROPHIC PATTERNS Hence, predators To introduce a bit more realism are ultimately lim- into my model, I varied the annual pri- ited by their prey mary productivity and the annual pre- W (Figure 2). dation rate on herbivores stochastically i Vegetation 5 Admittedly, (Figure 3a, 3b). Such stochastic varia- most predator-prey tion in primary productivity mimics 53 0 systems involve annual variation due to weather. I mod- 8 more than one prey elled the stochastic variation in avail- and one predator spe- ability of prey as a variation in wolf cies, yet some well- carrying capacity, which could result studied systems are from changes in snow conditions. For PRIMARY PRODUCTIVITY quite simple. On Isle parameter values that lead to realistic Royale in Lake Supe- relative densities of predator and prey, Figure l(a). Classic Top-Down Trophic Patterns. At very rior, for example, stochastic variation of primary produc- low primary productivity, vegetation can not support her- moose (Alces alces) tivity or predation rate, or both, affects bivores. When primary productivity is capable of support- have been basically all three trophic levels (Figure 3b). If ing herbivores, increased primary productivity leads to a the only prey for stochasticity in primary productivity is population of herbivores that maintains a stable standing wolves (Canis lu- superimposed on a long-term cycle, crop of vegetation. When primary productivity is high enough to support enough herbivores to support preda- pus) for nearly five which might be caused by cyclical tors, the predators maintain a stable population of herbi- decades (McLaren variation in climate, for example, the vores, and, therefore, the standing crop of vegetation in- and Peterson 1994). cycle may be seen in the predator and creases. When primary productivity is capable of sup- Similarly, muskoxen prey populations (Figure 3c). A re- porting a secondary carnivore, the population of primary carnivores remains constant, herbivore populations in- (Ovibus moschutus) searcher studying such a predator-prey crease, and the standing crop of vegetation is constant generally are prey system might conclude that interactions once again. Population sizes are not drawn to scale so only for wolves, or between predator and prey populations that carnivore populations can appear on the figure.

WHO CONTROLS WHOM? SIMULATED Model Predictions: TROPHIC PATTERNS *Predators reduce W *Across years, primary productivity sets equilibrium population sizes of both herbivores equilibrium $ and predators. population size LU of herbivores, 6 N ZG 5 1, a3 L? L? $5%Q cc----- Carnivores E32 t PRIMARY PRODUCTIVITY PRIMARY PRODUCTIVITY

Figure l(b). Simulated Top-Down Trophic Patterns mimic Figure 2. Who Limits Whom? Over long time spans, the classic pattern except that the graph of the herbi- primary productivity dictates when herbivores and car- vore population has a "hump" at levels of primary pro- nivores can maintain populations. Predators reduce ductivity that are capable only of supporting low carni- long-term populationsof herbivores below the level sup- vore population sizes. portable by primary productivity and allow consequent increases in the standing crop of vegetation.

Vol. 18 No. 4 2001 Endangered Species UPDATE 99 EQUILIBRIUM CONDITIONS STOCHASTIC VEGETATION and FOR PREDATOR-PREY , PREDATORS I 1 SIMULATIONS z 0 C !3# Vegetation 30 1 Herbivores 8a +.

U Predators Predalors J i- YEAR YEAR Figure 3(a). Sample population trajectories for the Figure 3(b). Trajectories for vegetation, herbivores and standing crop of vegetation and for population sizes predators using the same model parameters used in Fig- of herbivores and predators. ure 3a except that primary productivity and predation vary stochastically. Year-to-year variation in herbivore popu- lation size depends more on variations in primary pro- ductivity during the preceding two years than on varia- lead to population cycles when the chastic variation in tions in predation pressure. cycles are actually driven from below primary productiv- by variation in primary productivity. ity leads to a sirni- Consistent with the results of lar, though smaller, STOCHASTIC VEGETATION and Boyce and Anderson (1999), I found decrease in long- that variation in primary productivity term, average prey W had a profoundly greater effect on year- populations. Sto- N, V) to-year changes in herbivore popula- chastic variation in tions than did variation in predation both primary pro- 6 efficiency. The vast majority of the ductivity and preda- 53 year-to-year variation in the model her- tion, however, leads n bivore population could be explained to a modest (ca. 2%) B by the productivity of vegetation dur- increase in prey ing the preceding two years, while only populations, be- about a quarter could be explained by cause of the de- YEAR the variation in predation rates. On a creased predator year-to-year basis, the model predicts populations. Figure 3(c). Same as Flgure 3b except that a long-term that variation in food affects popula- Who controls cycle is imposed on primary productivity, such as might tion dynamics of herbivores more than whom, predators or be caused by climate cycles. The cyclic variation in pri- mary productivity caused cyclic variation in predator and does variation in predation (Figure 4). prey? The prey populations might be mistaken as cycles caused by Stochastic variation of predation predicts that each interactions between the population of predators and prey. rate leads to an approximate 10% de- controls the other crease in the long-term, average preda- but the control acts tor populations. This result is expected at different levels, or on different scales. (Figure 2). In contrast, on a year-to- for predators with a Type 3 functional Ultimately, predator populations are year basis, variation in productivity of response. Because predation rate in- controlled by the populations of their food causes more variation in herbivore creases in a concave-downward fash- prey, wYch &e, in &,dependent on population sizes than does variation in ion with high and increasing prey popu- the primary productivity of their foods. predation rates (Figure 4). lations, variations in prey population When primary productivity can support size below the average population size an herbivore population large enough Real predators and prey lead to larger changes in predation than to support carnivores, however, preda- Do real populations of predators do variation in prey populations above tion then decreases the long-term, av- and prey act as the model predicts, the average. For the same reason, sto- erage size of the population of its prey with control acting on two scales?

100 Endangered Species UPDATE Vol. 18 No. 4 2001 by the 1930s, and by and their predators, predators are ca- pable of limiting prey populations to WHO CONTROLS WHOM? the 1960s, porcu- levels below the levels they would Model Predictions: pine populations had grown to previ- reach without predation. In addition, ously unknown when prey populations do not sup- sizes. Fishers were port predators, they limit the abun- reintroduced in the dance of their food. 1960s, and thus I studied a growing Management implications population. Porcu- Reduction of a predator's population pines had two lim- size to increase the size of a prey popu- iting resources: win- lation is often considered by manage- TIME ter food and safe ment agencies (Gasaway et al. 1983). winter dens. During The models presented here indicate that Figure 4. Who Limits Whom? Annual variation in plants winter, porcupines reducing the population of a predator explains most of the variation in the herbivore popula- denned near their will have little effect on the year-to-year tion, while variation in predation rate explains consider- food trees. Ddg fluctuations in the populations of its ably less. the years of high prey but may affect the long-term mean population density, population size of prey. Figures 1, 2 Results from field studies support porcupines browsed hemlocks and and 3a illustrate how, when primary this hypothesis. white pines heavily during winters, of- productivity is high enough, predators The data for communities with ten stunting tree growth, Where small might limit prey from reaching the vertebrates in arctic and north tem- of hemlocks dwindled and died, population size the food supply can Perate ecosystems reviewed by porcupines abandoned the associated support. Consequently, reducing the Oksanen and Oksanen (2000) are dens. In my study area, most dens were population of a predator can be one op- consistent with control on two scales. hollow trees or hollow logs. ofl low tion to consider, with caution, when Similarly, data for communities with logs with holes at both ends were ac- faced with a threatened or endangered herbivorous or predatory and ceptable t~ porcupines before fishers population of a prey species. small, vertebrate predators reviewed were reintroduced (Brander 1973; Reduction of a predator population by Schmitz et al. (2000) are consis- Brander and Books 1973; Powell and must be considered with caution for tent with control on two scales. Brander 1977). As the popula- several reasons. First, Figure 3c illus- IvIcLaren and Peterson (1994) ar- tion grew, however, fishers killed por- trates that cyclic and stochastic fluctua- gued from data on wolves, moose and cupines in log dens with two entrance tions in primary productivity can lead balsam f~ (Abies balsamia) on Isle holes. A safe den near good winter food to large fluctuations in an herbivore Ro~ale,that predation by wolves con- became in short supply. Fishers caused population that might be confused with trolled the moose population. In addi- a significant decrease in the porcupine a predator-prey cycle. The models pre- tion, moose browsed their major win- population (Figure 5). sented here suggest that the year-to- ter food, balsam fir, so heavily that pro- Wehausen (19%) also documented year variation in abundance of herbi- duction of fr was limited when the a resurgent predator population taus- vores are caused more by fluctuations moose po~ulationswas high. Reanaly- ing the distinct decease of a formerly in food supply than by changes in pre- sis of their data yields results consis- healthy prey population, in this case dation. The same may be true of long- tent with model results: production of mountain lions ( concolor) taus- term cycles of prey populations. Sec- fir explains a significant amount of the ing the decrease of a mountain sheep ond, most predators prey on several year-to-year variation in the moose (Ovis canadensis) population. prey species, which, in turn, have more ~~ulation,while~ear-to-~earvdation Thus, field data support the hy- than one predator, yet the models pre- in the wolf population does not. pothesis that populations of real preda- sented here deal with single predator Istudiedfishers(Martes~ennanti) tors and prey exhibit controls on two and prey populations. The switching and their Prey, ~dcularl~porcupines scales. Most annual variation in popu- of prey by predators may prevent any (Erethizon dorsatum), in Michigan's lations of prey appears controlled by one predator hmlimiting any one prey Upper Peninsula in the 1970s (re- variation in food supplies, Yet, population (Murdoch and Oaten 1975), viewed by Powell 1993). Fishers had when primary productivity is high and the removal of one predator spe- been extir~atedintheu~perpeninsulaenough to support communities of prey cies may simply offer opportunity to

Vol. 18 No. 4 2001 Endangered Species UPDATE 101 Canadian Entomology 91:293-320. McLaren, B.E. and R.O. Peterson. 1994. PORCUPINE POPULATION Wolves, moose and tree rings on Isle TRENDS Royale. Science 266:1555-1558. Mech, L.D. 1992. Wolves of the High Arc- tic. Voyageur Press, Stillwater, Minnesota. 1200 Ottawa National Forest Upper Peninsula Michiga Mech, L.D. 2000. The Wolves of Minnesota: =Q 2000 Howl in the Heartland. Voyageur Press. Stillwater, Minnesota. 3 ::; Murdoch, W.W. 1966. Community structure, 3 V, 400 population control, and competition B a cri- 8 200 tique. American Naturalist 100:219-226. a 0 Murdoch, W.W. and A. Oaten. 1975. Preda- 1960 1965 1970 1975 1980 1985I tion and population stability. Advances in Ecological Research 9:1 - 125. YEAR Oksanen, L,S. D. Fretwell, J. Armda, and P. Niemela. 1981. Exploitation ecosystem in gradients of primary productivity. Ameri- Figure 5. Changes in the porcupine population on the can Naturalist 118:240-261. Ottawa National Forest. In 1962, porcupines lacked preda- Oksanen, L. and T. Oksanen. 2000. The logic tors, and the population density was extremely high. Fish- and realism of the hypothesis of exploita- ers were reintroduced In the mid-19608, and by the late tion ecosystems. American Naturalist 1970s porcupines were rare. Double bars for porcupine 155:703-723. populations in 1973-76 present data for both the study Polis, G.A. 1999. Why are parts of the world sites sampled in 1962 and 1970-73 and for an expanded green? Multiple factors control productiv- number of study sites. ity and the distribution of green biomass. Oikos 86:3-15. Polis, G.A. and D.R. Strong. 1996. Food web complexity and community dynamics. American Naturalist 147:813-846. another. Finally, long-term control of cian 5(4):425-433. Powell, R.A. 1993. The fisher: Life history, a predator population when primary Brander, R.B. and D.J. Books. 1973. Return ecology and behavior. 2ndedition. Univer- of the fisher. Natural History 82(1):52-57. productivity varies stochastically may sity of Minnesota Press, Minneapolis. Elton, C. 1927. Animal Ecology. Sedgwick Powell, R.A. and R.B. Brander. 1977. Adap- lead to extirpation of the predator, es- & Jackson, London. tations of fishers and porcupines to their pecially when the prey is also subject Fretwell, S.D. 1977. The regulation of plant predator-prey system. Pp 45-53 in R.L. to control, as through harvest. If long- communities by food chains exploiting Phillips and C. Jonkel, eds. Proceedings term harvest of a prey species exceeds them. Perspectives in Biology and Medi- of the 1975 Predator Symposium. Mon- cine 20: 169-185. of its equilibrium population size, tana Forest & Conservation Experiment 10% Fretwell, S.D. 1987. Food chain dynamics: Station, University of , Missoula. and primary productivity fluctuates as The central theory of ecology? Oikos Schmitz, O.J., P.A. Hamback and A.P. in my models, any control of the preda- 50:291-301. Beckerman. 2000. Trophic cascades in ter- tor population could lead to its extirpa- Gasaway, W.C., R.O. Stephenson, J.L. Davis, restrial systems: A review of the effects of P.E.K. Shepard, and O.E. Burris. 1983. In- tion within decades. carnivore removals on plants. American terrelationships of wolves, prey, and man Naturalist 155: 141- 153. in interior Alaska. Wildlife Monographs Strong, D.R. 1992. Are trophic cascades all Literature cited 84: 1-50. wet? Differentiation and donor-control in Boyce, M.S. andE.M.Anderson. 1999. Evalu- Gray, D. 1987. The muskoxen of Polar Bear speciose ecosystems. Ecology 73:747-754. ating the role of carnivores in the greater Pass. Fitshenry & Whiteside, Markham, Wehausen, J.D. 1996. Effects of mountain Yellowstone ecosystem. 4 265-284 in TOW. Ontario. lion predation on bighorn sheep in the Si- Clark, E.P. Curlee, S.C. Minta, and P.M. Hairston, N.G., F.E. Smith, and L.B. erra Nevada and Granite Mountains of Kareiva, eds. Carnivores in ecosystems: The Slobodkin. 1960. Community structure, California. Wildlife Society Bulletin Yellowstone experience. Yale University population, and competition. American 24:471-479. Press, New Haven, Connecticut. Naturalist 94:42 1-425. White, T.C.R. 1978. The importance of a Brander, R.B. 1973. Life history notes on the Holling, C.S. 1959. The components of preda- relative shortage of food in animal ecol- porcupine in a hardwood-hemlock forest tion as revealed by a study of small mam- ogy. Oecologia 3:71-86. in Upper Michigan. Michigan Academi- mal predation of the European pine sawfly.

102 Endangered Species UPDATE Vol. 18 No. 4 2001 A New Approach to Understanding Canid Populations Using an lndividual-based Computer Model: Preliminary Results

William C. Pitt USDNAPHISMIS, National Wildlife Research Center, Utah State University, Logan, UT, Utah State University, Logan, Utah, 84322-5295;(435) 245-6091 ; wwQcc.usu.edu Frederick F, Knowlton USDNAPHISMIS, National Wildlife Research Center, Utah State University, Logan, Utah, 84322-5295 Paul W. Box Department of Geography and Earth Resources Department, Utah State University, Logan, Utah, 84322

Abstract Ensuring the welfare of wild canid populations depends upon the ability to integrate species biology, the environmental aspects upon which those populations depend, and the factors control- ling species abundance. Toward this end, we developed an individual-based computer model using Swarm to mimic natural populations. Swarm is a sofhvare platform that allows the user to describe individual behaviors for all individuals, link those behaviors in each concurrent time step, and assemble behaviors and objects in a hierarchicalfvamework. Our model stands apartfvom previous modeling eforts because it relies on field data and explicitly incorporates behavioral features, such as dominance and territoriality, as major determinates of species demography. Individual variation, such as status within territorial social groups and age-based reproduction are assumed, but assumptions typically associated with most demographic models are not needed. The eventual goal is to incorporate other environmental components such as prey abundance andor competing carnivores. This type of model could also provide insights into potential management alternatives for when the gray wolf is removed from endangered status in Minnesota.

Introduction clude the individual characteristics The model Ensuring the welfare of wild canid that are critical to canid populations Swarm is a software platform that populations depends upon the abil- and past computer models have not allows the user to describe indi- ity to integrate our best understand- incorporated territoriality and so- vidual behaviors for all individuals, ings of species biology, the environ- cial structure (Connolly and link those behaviors in each con- mental aspects upon which those Longhurst 1975). Toward this end, current time step, and assemble be- populations depend, and the factors we developed an individual-based haviors and objects in a hierarchi- controlling species abundance computer model using the Swarm cal framework (Savage and (Gese et a1 1989; Murray et a1 modeling system to provide a bet- Askenazi 1998; Railsback et al. 1999). Previously, biologists and ter understanding of canid popula- 1999; SDG 2001). Our model managers have relied upon insights tion dynamics. We use stands apart from previous model- provided by many analytical and (Canis latrans) to parameterize the ing efforts because it relies on field computer models of animal popu- model for this exercise, but the data with all population parameters lations. Canid populations, how- model could easily be adapted to derived from data sets and pub- ever, differ from other species be- many other canid species with simi- lished papers, and explicitly incor- cause they are highly territorial and lar population structure. This paper porates behavioral features, such as have a specific social structure, is a preliminary summary of a dominance and territoriality, as relatively low density (Knowlton model that will be presented in major determinates of species de- 1972; Sillero-Zubiri and Gotelli greater detail elsewhere (Pitt et al. mography (Connolly and Longhurst 1995; Knowlton et. a1 1999). Ana- in preparation) 1975; Knowlton et al. 1999; Pitt et lytical models are not suited to in- al. 2001). Individual variation,

Vol. 18 No. 4 2001 Endangered Species UPDATE 103 such as status within territorial social sity (Crabtree and Sheldon 1999; Model output groups and age-based reproduction Knowlton et al. 1999). Field evi- We ran the population model under are specified and assumptions typi- dence for and against this argument three management scenarios: no re- cally associated with most demo- has been mixed (Gier 1968; Todd moval, pulse removal (a proportion graphic models are not needed. et al. 1981 ; Knowlton and Stoddart of animals were removed in year The coyote population model 1983; Windberg 1995; Gese et al. five and the population then al- was divided into 100 packs and a 1996). The most likely reason for lowed to recover), or press removal collection of non-territorial ani- these mixed results is that the num- (a constant proportion of animals mals. Each individual is character- ber of offspring produced is a func- were removed every year after year ized by sex, age, status, and pack tion of the food supply for that par- five). The populations were evalu- membership. Pack size was not lim- ticular female. The food supply is ated according to structure, the re- ited but the likelihood of subordi- a function of the food in the terri- sistance to removal (proportion of nates increased with the number of tory and the number of animals in animals removed required to have animals in the pack. Individuals the pack. Studies that have at- an effect for more than one year), could change status or pack mem- tempted to determine the relation- and the resilience of the population bership by dispersing from natal ship between offspring produced (how quickly the population recov- packs, replacing a dominant animal, and density of food supply have ered under various removal levels). or by moving to a pack from non- looked at entire populations and With no removal, the popula- territorial status. large land areas (Gier 1968; Todd tion was stable and population size As with most animals, the prob- et al. 1981; Knowlton and Stoddart ranged from 350 to 700 adult ani- ability of mortality increases with 1983; Windberg 1995; Gese et al. mals with 15 to 35% of the popula- age. Mortality rates are higher for 1996). Thus, the relationship tion being non-territorial. The rea- non-territorial animals than pack would only be observed if most son for this stability was animals members (Gese et al. 1989). In ad- packs were similar in size, food were forced out of packs as they dition, mortality rates increase with supply was homogenous across the matured and non-territorial animals the density of non-territorial animals landscape, and territories were had a higher mortality rate then because they would potentially share identical in size. The likelihood of animals in packs. The population a common area and the probability all of these factors being similar in exhibited source-sink dynamics. of encountering other animals would one population would be low and Average pack size in this simula- increase with density. Thus, density extremely rare between popula- tion was about four but varied from increases would either result in less tions, so this relationship would not one to eight. food or an increase in the number of be observed under most conditions To determine the effect of pulse negative encounters. on a population basis. For these removal on the population, we let Although subordinate coyotes reasons, we set the mean number of the population run for five years occasionally produce offspring in pups produced as a function of the and then randomly removed 10 to natural populations, in our model number of pack members. In this 90% of the adult population in one only alpha females breed model, territories are identical so year and then examined the re- (Knowlton et al. 1999). The birth we could ignore differences be- sponse of the population. All popu- rate is based on a normal distribu- tween territories. lations recovered within one year tion with the mean based upon pack The second part of this model- when less than 60% of the popula- size. Few would disagree that the ing exercise is the management tion was removed (Figure 1). Ba- number of offspring produced is a model, which allows us to examine sically, the population was reduced function of the health of the animal. the effects of managing the popu- until new offspring were produced. There has been, however, continued lation (Pitt et al. 2000). The man- The number of transients decreased disagreement over what is a good agement model combines the popu- as animals moved into packs and indicator of health. Some evidence lation model and a manipulation fewer animals moved out of packs. from captive coyote studies sug- model so we can investigate the ef- Populations subject to removal had gests that old (>8 years) animals fect of removing individuals on younger age structures. When more will produce fewer offspring population size and the resistance than 60% of the population was re- (Green et al. 2001). The most con- and resilience of the population. moved, the population required tentious argument is that litter size Herein, only random removal indi- more than one year to return to the is a function of food supply or den- viduals will be considered. population size prior to removal.

104 Endangered Species UPDATE Vol. 18 No. 4 2001 population decline could take sev- eral more years because territories remained in the model and animals did not move to mate, natural mor- tality rates were not reduced at low densities, and animals did not move in from surrounding areas. Remov- ing more than 70% of the popula- tion annually would become logis- tically difficult at low densities. Territoriality would likely dissolve at low densities, animals would move to mate, natural mortality may be re- duced, animals would immigrate into the population, and the high removal rates could not be achieved.

Implications for management These simulations suggest that coy- otes and other canid populations are very resistant and resilient to I0 20 30 40 50 60 70 80 90 change. A population decrease was Proportion Removed not observed until more than 60% of the population was removed an- Figure 1. Number of years required for a population to return to pre-removal nually. The populations are buff- size after a certain proportion of animals are randomly removed from the popu- ered against change by the high re- lation in one year. The horizontal line indicates the threshold where annual productive capacity and the non- reproduction compensates for animals removed. breeding animals in the population, subordinates and non-territorial animals. Non-breeders would re- This removal proportion is lower population, we let the population place breeding individuals lost from than reported in previous computer run for five years and then ran- the population so the reproductive models (Connolly and Longhurst domly removed 10 to 90% of the capacity of the population is not re- 1975). The population recovered adult population each year and ex- duced. Coyote and other canid within five years, however, even amined the response of the popula- populations are resilient because with 90% removal in one year. tion. When removal was less than they have a high reproductive ca- When a large (>70%) proportion of 60% of the population, population pacity. These conclusions may pro- the population was removed, the size was the same as an unexploited vide insight into the potential ef- number of non-territorial animals population, and it did not decline, fects of disease on Ethiopian decreased. In natural populations even after 50 years of simulation. wolves (Canis simensis) or the po- less time would be required to re- The population structure, however, tential management of timber cover than what was depicted in the differed from an unexploited popu- wolves (Canis lupus). These spe- model because in the model, terri- lation. The population with press cies would also have similar popu- tories remained even at low densities, removal at 50% had fewer transient lation characteristics as was dis- animals were not allowed to move animals (10 to 25%), a younger age played in this model. out of their territories to mate, and structure, and higher reproduction In the future, the model analy- animals were not allowed to move in than an unexploited population. sis will be expanded to investigate from surrounding areas. Further- High removal rates (>70% per year) various types of removal, e.g. se- more, we did not reduce natural mor- resulted in an initial loss of non-ter- lective versus random removal. tality rates at low densities. ritorial animals and after seven The management model allows the To determine the effect of sus- years the entire population was re- user to remove animals of a specific tained or press removal on the moved. In natural populations, a status, litters of offspring, during a

Vol. 18 No. 4 2001 Endangered Species UPDATE 105 particular month, and/or animals in Gier, J.T. 1968. Coyotes of , Kan- of management on territorial animals a specific area. In addition, we can sas Agricultural Experiment Station Bul- using Swarm. Pp 36-40 in W. C. Pitt, letin 393, l l8pp. ed. Swarm User Group Conference Pro- also investigate the effects of repro- Green, J.S., F.F. Knowlton, and W.C. Pitt. ceedings. Published by S. J. and Jessie ductive control or the effects of dis- 2001. Reproductive patterns among cap- E. Quinney Natural Resources Research ease on populations. To determine tive wild-caught coyotes (Canis latrans). Library, Logan, UT. the effects of unequal resource dis- Journal of Mammalogy in press. Pitt, W.C., F.F. Knowlton, and P.W. Box. tribution, we can also vary re- Knowlton, F.F. 1972. Preliminary interpreta- In preparation. A new approach to un- tions of coyote population mechanics with derstanding canid populations using an sources among packs and over time some management implications. Journal of individual-based computer model. as part of the foraging and preda- Wildlife Management 36369-382. Railsback, S.F., R.H. Lamberson, B.C. tion models. Other components we Knowlton, F.F. and L.C. Stoddart. 1983. Harvey, and W.E. Duffy. 1999. Move- contemplate adding are competing Coyote population mechanics: another ment rules for individual-based models look. Pp 93-1 11 in F.L. Bunnell, D.S. of stream fish. Ecological Modelling carnivores, predator-prey interac- Eastman, and J.M. Peek, eds. Sympo- 123:73-89. tion, as well as, management cost- sium on Natural Regulation of Wildlife Savage, M. and M. Askenazi. 1998. benefit programs to the model. Populations. Forest, Wildlife and Range Arborscapes: a Swarm-based multi- Experiment Station, University of Idaho, agent ecological disturbance model. Literature cited Moscow, Idaho. Proceedings of Geographical and Envi- Knowlton, F.F., E.M. Gese, and M.M. Jae- ronmental Modelling, Available as Santa Connolly, G.E. and W.M. Longhurst. 1975. ger. 1999. Coyote depredation control: Fe Institute Working Paper 98-06-056. The effects of control on coyote popu- an interface between biology and man- Sillero-Zubiri, C. and D. Gotelli. 1995. lations: a simulation model. Division of agement. Journal of Range Management Spatial organization in the Ethiopian Agriculture Science Bulletin 1872, 52:398-412. wolves Canis simensis: large packs and Davis, CA. Murray, D.L., C.A. Kapke, J.J. Everman, small stable home ranges. Journal of Crabtree, R.L. and J.W. Sheldon. 1999. and T.K. Fuller. 1999. Infectious dis- Zoology (London) 237:65-8 1. Coyotes and canid coexistence in eases and the conservation of free-rang- SDG (Swarm Development Group). 2001. Yellowstone. Pp 127-163 in T. W. Clark, ing large carnivores. Animal Conserva- The Swarm Simulation System. Pp 1- A. P. Curlee, S. C. Minta, and P. M. tion 2:241-254. 10 in W.C. Pitt, ed. Swarm 2000 Con- Karieva, eds. Carnivores in Ecosystems, Pitt, W.C., F.F. Knowlton, A. Ogawa, and ference Proceedings. Published by S. J. The Yellowstone Experience. Yale Uni- P.W. Box. 2000. Evaluation of depreda- and Jessie E. Quinney Natural Resources versity Press, New Haven. Connecticut. tion management techniques for territo- Research Library, Logan, UT. Gese, E. M., 0. J. Rongstad, and W. R. rial animals using a computer model: Todd, A.W., L.B. Keith, and C.A. Fisher. Mytton. 1989. Population dynamics of coyotes as a case study. Pp 315-318 in 1981. Population ecology of coyotes dur- coyotes in southeastern Colorado. Jour- T.P. Salmon and A.C. Crabb, eds. Pro- ing a fluctuation of snowshoe hares. Jour- nal of Wildlife Management 53: 174- 181. ceedings of the 191hVertebrate Pest Con- nal of Wildlife Management 45629-640. Gese, E.M., R.L. Ruff, and R.L. Crabtree. trol Conference. Published by Univer- Windberg, L.A. 1995. Demography of a 1996. Social and nutritional factors in- sity of California, Davis. high-density coyote population. Cana- fluencing the dispersal of resident coy- Pitt, W.C., A. Ogawa, F.F. Knowlton, and dian Journal of Zoology 73:942-954. otes. Animal Behaviour 52:1025-1043. P.W. Box. 2001. Evaluating the effects

106 Endangered Species UPDATE Vol. 18 No. 4 2001 Habitat issues Mapping Occupied Habitat for Forest Carnivores in the American West and Estimating their Conservation Status

David L, Gaillard Predator ConservationAlliance, P.O. Box 6733,Bozeman, MT 59771;(406) 587-3389; [email protected];www.predatorconservation.org

Abstract Conservation concerns are escalating due to the small numbers, reduced range, and increasingly fragmented distribution of wide-ranging forest carnivores in the American West-speci$cally the (Lynx canadensis), (Gulo gulo), andfisher (Martes pennanti). Observation data from the U.S. Forest Service and state natural heritage programs were compiled and mapped. Occupied habitats and population centers for the three species were identijied and population sizes estimated based upon recorded densities and distribution. The results indicate small, isolated populations well below what may be necessary for long-tern viability.

Introduction they are limited to specific study fisher based on available presence1 Some of the smaller forest carni- areas and fail to address the con- absence data, and to estimate popu- vores of the American West-the servation status of each species lation sizes within these areas based lynx (Lynx canadensis), wolverine throughout its range. upon recorded densities, as a first step (Gulo gulo), and fisher (Martes We can no longer take for toward devising a long-term conser- pennunti)-have not yet grabbed granted the survival of wide-rang- vation strategy for these species. headlines like (Ursus spp.) ing, low-density forest carnivores. and wolves (Canis lupus). Yet As we have done for the grizzly Mapping occupied habitat mounting concerns about their vi- bear and the wolf, we should make To identify occupied habitats and ability indicate that large carnivores conscious decisions about where in population centers for the lynx, are not the only species in jeopardy the western U.S. we will maintain wolverine, and fisher, observation because of past and ongoing habi- and restore forest carnivores, and data from the U.S. Forest Service tat loss and fragmentation, and ex- protect them and their habitat in and the natural heritage programs cessive human-caused mortality. these areas accordingly. The capac- of eight western states were gath- While these lesser known species ity of these areas to support forest ered (Maj and Garton 1994; were never persecuted like the wolf, carnivores should be assessed to McKelvey et al. 2000b). These data grizzly bear (Ursus arctos), and ensure that sufficient habitat is pro- were not collected through stan- other species that pose a risk to live- tected to provide for their long-term dardized survey techniques but in- stock, their conservation status is survival and recovery. Given the clude observations made by work- now suffering due to neglect. forest carnivores' large ranges and ers in the field, trapping records, Critical information about the low densities and the fragmented and museum specimen records. habitat requirements and popula- nature of suitable habitat that re- State natural heritage programs tion sizes of these animals is lack- mains, a conservation strategy will maintain this data and attempt to ing, leaving conservationists ill-in- likely require restoration and main- filter out unreliable observations. formed at where and how to initiate tenance of a connected network of GIs software was used to high- conservation strategies. Existing population centers across the west- light parcels of public lands where range maps are too coarse to pri- ern U.S. and (e.g., the observations were located. Pri- oritize specific areas for protection, McKelvey et al. 2000a). The ob- vate land was excluded from this such as individual mountain ranges jective of this paper is to identify analysis, because of the difficulty or watersheds. There are few pub- occupied habitats and population delineating borders around point lished population estimates, and centers for the lynx, wolverine, and locations on private lands. Admin-

Vol. 18 No. 42001 Endangered Species UPDATE 107 istrative boundaries were conve- prises 90 to 100% of these nient for this purpose on public areas. Copeland (1996) es- lands, and though they have no bio- timated one wolverine per logical basis they serve the purpose 90 to 248 km2 within his of delineating the large landscape study area in the Sawtooth features relevant to this scale of Mountains, based upon his analysis. This methodology unfor- analysis of 1,050 reloca- tunately excludes important areas tions of 19 by of carnivore habitat on private ground and aerial telem- lands, but the vast majority (>80%) etry. A conservative esti- of the carnivore observations oc- mate of one wolverine per curred on public lands. 200 km2 across each of These areas were sorted by date these areas indicates three Figure 1. Occupied habitat for lynx, wolverine, to provide an approximation of cur- subpopulations of 100 wol- and rent versus historic range. Observa- verines each in Idaho (Fig- tions made after 1990 were used to ure 2). Delineation of po- delineate current range while obser- tential subpopulations for Dark Cny = hbSc and vations prior to 1990 denote historic each of the forest carni- tnbl kndr wiIh cumnl range for the purpose of this analy- vores and estimating their Obdt,VP11W111WPl LighlGmy = Fuhlte and WlwiS*iibpari sis. Overlapping data for all three numbers provides a starting

108 Endangered Species UPDATE Vol. 18 No. 4 2001 estimated numbers of lynx est carnivore observation data free of (Figure 4) and grizzly bear charge. Thanks also to the Ecology across the western U.S., both Center in Missoula, Montana for of which are listed as threat- some GIs assistance, to Defenders of ened species under the fed- Wildlife for providing a venue to eral Endangered Species Act. share this analysis, and to Stephen Kendrot (USDA Wildlife Services) Discussion for a helpful review. This analysis provides a coarse but useful first step Literature cited toward identifying occupied Copeland, J. 1996. Biology of the wolverine in habitat for the lynx, wolver- central Idaho. MS Thesis, University of Idaho, Moscow, Idaho, May 1996. ine, and fisher, toward as- Groves, C.R. 1988. Distribution of the wolver- Figure 3. Fisher sub-population estimates sessing their conservation ine in Idaho as determined by mail question- status across the American naire. Northwest Science 62(4): 181-5. West and prioritizing areas Maj, Mary and E.O. Garton. 1994. Fisher, lynx, for their protection. Obser- wolverine; summary of distributiorl informa- tion. Pp 169-175 in L.F. Ruggiero, K.B. vation data are inherently Aubry, S.W. Buskirk, L.J. Lyon, and W.J. biased by observer effort Zielinski, eds. The scientific basis for con- and reliability, and by im- serving forest carnivores: American , perfect reporting and orga- fisher, lynx, and wolverine in the western United States. U.S. Dept, ofAgriculture,For- nization of observations. est Service, Gen. Tech. Rep. RM-254. These data should be re- McKelvey, K.S ., K.B. Aubry, J.K. Agee, S. W. fined with more careful Buskirk, L.F. Ruggiero, and G.M. Koehler. 2000a. Lynx conservation in an ecosystem 1)rrk Gray = Public and tnbal lands screening and supple- wrlh cumnloh\ew;llionr (IRLl's) mented with new informa- management context. Chapter 15 in Ltgh~Gray = Publte and tribal Ian&% Ruggiero, L.F., K.B. Aubry, S.W. Buskirk, with paslohcwauons (prs I9Ws) tion regarding the conser- G.M. Koehler, C.J. Krebs, K.S. McKelvey, vation status and distribu- and J.R. Squires, eds. Ecology and conser- tion of these species. Addi- vation of lynx in the United States. U.S. Figure 4. Lynx sub-population estimates tional data from Canada are Dept. of Agriculture, Forest Senrice Rocky Mountain Research Station,Missoula, Mon- needed to develop a com- tana; published by University Press of Colo- prehensive, range-wide rado, Boulder, Colorado. 1987), and that just a fraction of the conservation assessment and strat- McKelvey, K.S ., K. B. Aubry, andY.K. Ortega. individuals within these carnivore egy for the forest carnivores. De- 2000b. History and distribution of lynx in populations are actively breeding spite these limitations, this analy- the contiguous United States. Chapter 8 in Ruggiero, L.F., K.B. Aubry, S.W. Buskirk, and thus count toward the effective sis provides an initial step toward G.M. Koehler, C.J. Krebs, K.S. McKelvey, population size (Ruggiero et al. the design and implementation of a and J.R. Squires, eds. Ecology and conser- 1994). The fisher appears to be the forest carnivore conservation strat- vation of lynx in the United States. U.S. most imperiled of the three in the egy that encompasses occupied Dept, of Agriculture, Forest Service Rocky Mountain Research Station, Missoula, Mon- West, with an estimated total popu- habitat within the western U.S. for tana; published by University Press of Colo- lation of perhaps 600 animals, includ- one or more of these species. rado, Boulder, Colorado. ing populations that may be function- Ruggiero, L.F., K.B. Aubry, S.W. Buskirk, L.J. ally isolated in the southern Sierra, Acknowledgements Lyon, and W.J. Zielinski, eds. 1994. The northern Sierra, southwestern Or- I am grateful to the U.S. Forest Ser- scientific basis for conserving forest cami- vores: American marten, fisher, lynx, and egon, and portions of Greater vice Rocky Mountain Research Sta- wolverine in the western United States. Gen Yellowstone (Figure 3). The wolver- tion for compiling and providing Tech Rep RM-254, Fort Collins, Colorado, ine is more widespread than the me with its lynx distribution data, U.S. Dept. of Agriculture, Forest Service, fisher, but its low densities coupled and to the natural heritage programs Rocky Mountain Forest and Range Exp. Stat, 184 pp. with recent declines in the Sierra and of California, Colorado, Idaho, Soul6 M.E. 1987.Viable populations for con- Northwest are cause for concern. Montana, Oregon, Utah, Washing- servation. Cambridge University Press, Population estimates for both the ton, and for providing for- Cambridge, England. fisher and the wolverine are less than

Vol. 18 No. 4 2001 Endangered Species UPDATE 109 Road Density as a Factor in Habitat Selection by Wolves and Other Carnivores in the Great Lakes Region

Adrian P, Wydeven Mammalian Ecologist/ConservationBiologist, Wisconsin Department of Natural Resources, 875 South 4th Ave., Box 220, Park Falls, WI 54552; (715) 762.4684 x107; (fax) (715) 762-4348; [email protected] David J. Mladenoff Department of Forest Ecology and Management; University of Wisconsin Madison; 1630 Linden Drive; Madison, WI 53706 Theodore A, Sickley Department of Forest Ecology and Management; University of Wisconsin Madison; 1630 Linden Drive; Madison, WI 53706 Bruce E. Kohn Wisconsin Department of Natural Resources; Highway 17 S., Box 576; Rhinelander, WI 54501 R. P. Thiel Wisconsin Department of Natural Resources; Sandhill Wildlife Area; Box 156; Babcock, WI 54413 Jennifer L. Hansen Land Information and Computer Graphics Facility; University of Wisconsin Madison; 550 Babcock Drive, 8102 Steenbach Library; Madison , WI 53706; (608) 263-5534; (fax) (608) 262-2500; [email protected]

Abstract Although wolves (Canis lupus) and many other carnivores are habitat generalists, certain land- scape features can be used to predict suitable habitat. Thiel examined the concept of road density as an important factor in the persistence of wolfpopulations in Wisconsin prior to the 1960s and found a relationship with the disappearance of breeding wolfpopulations when average road density exceeded 0.58 km/km2. Mladenofand co2leagues examined road density in the early 1990s as a factor in predicting favorable habitat of wolves colonizing Wisconsin between 1980 and 1992, and found that areas with road densities less than 0.45 km/km2had greater than a 50% probability of being colonized by wolf packs. Mladenoff and colleagues updated this work in the late 1990s by examining 23packs colonizing Wisconsin between 1993 and 1997; 78% continued to occupy areas with road densities below 0.45 km/km2. In a recent examination of radio-collared wolves in Wisconsin, a total of 60% of human-induced mortality occurred at road densities above-0.63 km/km2. Although road density may become less of a factor as human tolerance changes, and wolfpopulations increase, it continues to be an importantfactor in habitat selection by wolves and probably other carnivores.

Introduction 1992; Jensen et al. 1986; Mech et wolves were extirpated from Wis- Gray wolves (Canis lupus) are gen- al. 1988; Mladenoff et al. 1995; consin by 1960 (Thiel 1985; eralists in their use of habitat, and Thiel 1985). Wydeven et al. 1995). historically have been found in Thiel(1985) examined the dis- most regions across temperate, bo- Early development of concept appearance of breeding populations real, and arctic regions of North of road density of wolves in Wisconsin from 1926 America (Mech 1995). Despite this As a graduate student under Aldo to 1960. Using State Highway generalist nature of habitat use, Leopold, Thompson (1952) studied Commission Reports, he deter- landscape features, especially those wolf food habits in northern Wis- mined that breeding wolves disap- relating to human impacts, can be consin in the late 1940s, about 10 peared from 10 Wisconsin counties used to predict suitable wolf habi- years before wolves disappeared after road densities in these coun- tat (Corsi et al. 1999; Massolo and from the state. Thompson (1952) ties exceeded 0.48 to 0.68 krn/km2 Meriggi 1998; Mladenoff et al. warned that development and open- (X=0.58 km/km2). The value of 0.6 1995). Road density has frequently ing of roads across the logged for- krn/km2 has since been used fre- been used as a landscape feature to ests of northern Wisconsin could quently as the threshold level at predict suitable wolf habitat (Corsi cause wolves to become extirpated which wolf populations can be et al. 1999; Frair 1999; Fuller et al. from the state. As predicted, maintained. This level was found

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110 Endangered Species UPDATE Val. 18 No. 4 2001 to correspond well with areas of oc- pared to 14 random non-pack areas Mladenoff et al. (1995) used cupied wolf range in Minnesota scattered across northern Wiscon- road density as the main factor used (Fuller et al. 1992; Mech et al. 1988), sin. Areas occupied by wolf packs in a logistical regression model that Michigan, and Ontario (Buss and (80% isopleth of harmonic mean) predicted areas of suitable wolf Almeida 1998; Jensen et al. 1986). had average road densities of 0.23 habitat (Table 2). Areas with < 0.45 kmlkm2 (Table 1). Road density km/km2 were considered highly GIs analysis of road density was based on paved roads, and im- suitable wolf habitat and were esti- Through elimination of bounties proved dirt and gravel roads that mated to have > 0.50 probability of and protection by the 1973 Endan- appeared as solid lines on U.S. Geo- being colonized by wolf packs. gered Species Act, wolves were logical Survey (USGS) 1: 100,000 Minnesota had the most extensive provided protection that allowed re- quadrangle maps (Mladenoff et al. area of highly suitable habitat, and 'colonization of Wisconsin in the 1995). Other important features of packs expanded outside perceived 1970s (Wydeven et al. 1995). wolf pack areas included lack of suitable habitat in some areas (Berg Mladenoff et al. (1995) used a geo- urban and agricultural areas, exten- and Benson 1998). Fuller et al. graphic information system (GIs) sive forest (X=93%), high percent- (1992) indicated that in 1989, 88% to assess landscape features that age wetland (X=29%), mostly pub- of wolf pack areas had road densi- contributed to re-colonization of 14 lic lands and industrial forest land ties < 0.70 krn/km2. As predicted, Wisconsin wolf packs from 1980 to ( X=80%), and low human popula- most areas of highly suitable habi- 1992. Known pack territories of tion density. Human density com- tat were occupied by wolves in radio-collared wolves were com- pared closely to road density. Michigan (James Hammill, per- sonal communication). Based on the logistical regres- Table 1. Average landscape variables for 14 wolf packs, random non- sion model, areas with > 0.60 km pack areas (1-144)and overall Wisconsin study area (modified from road/km2 have less than a 10% Mladenoff et al. 1995). chance of being occupied by wolf packs. Thus, the GIs analysis agrees with the threshold found by Thiel(1985).

Roads as a factor in wolf habitat in other studies A habitat model developed in the Great Lakes region was used as a basis for assessing potential wolf habitat in the northeast U.S. (Harrison and Chapin 1998; Mladenoff and Sickley 1998). Mladenoff and Sickley (1998) esti- mated 53,500 km2of potential habi- tat in Maine and New Hampshire and 16,020 km2 in New York. Harrison and Chapin (1998) esti- mated 48,787 km2of potential habi- tat in Maine and New Hampshire, and 14,618 km2 in New York. Mladenoff and Sickley (1998) re- lied mainly on road density values, while Harrison and Chapin used a combination of road density and human population density. Road density has also been found to be important in predicting

Vol. 18 No. 4 2001 Endangered Species UPDATE 111 Table 2. Area of wolf habitat probability classes from a logistical curring in areas that seemed unsuit- regression model and corresponding road density for portions of able (road density > 0.60 km/km2), three Great Lakes states (from Mladenoff et el. 1995). were within 10 km of areas of highly suitable habitat. We recently examined the rela- tionship between wolf mortality and road density. Fifty radio-col- lared wolves died in Wisconsin be- tween 1979 and 1999. The road densities for 47 wolf mortalities were collected (Figure 1). The av- erage road density for natural mor- talities was 0.65 km/km2 and for human-induced mortalities was 0.78 km/km2. Highest natural mortality was at wolf habitat in Italy (Corsi et al. 1997. Five packs (22%) exceeded the road density range of 0.63 to 1999; Massolo and Meriggi 1998). the 0.45 km/km2 threshold of road 0.84 km/km2, areas considered poor The estimated area of highly suitable density and one (4%) exceeded the habitat. We initially had expected habitat in Italy (14,200 krn2) (Corsi 0.60 km/km2 threshold. Thirteen natural mortality to be highest in et al. 1999) was similar to the area packs were radio collared and pro- areas of most suitable habitat where estimated in Wisconsin (14,864 km2) vided more precise data on area of wolves most frequently occurred, (Mladenoff et al. 1995). pack use; two (15%) exceeded the but higher rates at higher road den- Kohn et al. (2000) conducted a 0.45 km/km2 threshold and one sity make sense. Wolves dying variety of studies examining the re- (7%) exceeded the 0.60 km/km2 from natural mortality, died mainly lationship of wolves to roads in threshold. In general, the road den- from disease or intraspecific strife. northwest Wisconsin. Shelley and sity model continued to be a good Diseased animals lose their fear of Anderson (1995) found road den- predictor of suitable wolf habitat. humans, and often wander off by sities in northwest Wisconsin wolf The one that exceeded the themselves into poor habitat. An territories to average 0.33 km/km2 0.60 krn/km2threshold was in a state adult female with severe mange for eight pack areas. Although wildlife area that had a higher road crawled into a garage in 1993. wolves selected areas of low road density (0.71 km/km2), but greater Mortality from intraspecific strife density, travel areas selected by access control may have nullified the usually occurs near the edge of a packs were generally close to trails effects of higher road density. territory and often pack boundaries and forest roads (Gehring 1995). The wolf population in Wiscon- are near roadways. Thus even Unger (1999) found that wolves se- sin increased from 15 wolves in wolves dying from natural mortal- lected den sites in roadless or low 1985 to 248 wolves in 2000, total- ity are more likely to be killed road density areas; dens were gen- ing 66 packs (Wydeven and closer to roads. erally located more than 1 km from Wiedenhoeft 2000). We examined Human-induced mortality improved roads. Frair (1999) found whether packs occurred within ar- peaked at relatively high road den- that wolves most frequently used eas of suitable habitat as illustrated sities (Figure 2). Most shootings areas away from roads, and aver- by Mladenoff et al. (1995, 1999). and vehicle collisions occurred at age road density in wolf territories Of 66 packs in Wisconsin, 53 (80%) road densities of 0.84 to 1.14 km/ was 0.25 km/km2;road density was were contained in areas mapped as km2. A total of 60% of human-in- found to be the best predictor of highly suitable wolf habitat (P > duced mortality occurred at a road suitable wolf habitat. 0.50), seven (1 1%) were contained density greater than 0.63 krn/km2. within marginal wolf habitat (0.50 In general, wolves appear more Recent examinations of road > P > 0.10), and six (9%) occurred likely to be killed at higher road densities in Wisconsin in areas mapped as poor wolf habi- densities. Although most wolves Mladenoff et al. (1999) examined tat (P < 0.10). Thus, packs contin- spend little time at these higher 23 additional wolf territories that ued to occur mainly in areas of low densities, they are at a much greater colonized Wisconsin from 1993 to road density. Even those packs oc- risk of being killed in these areas.

1 12 Endangered Species UPDATE Vol. 18 No. 4 2001 9 certain types of roads and seem more attracted to areas with low 8 traffic volume (Lovallo and 7 Anderson 1996). American mar- ten (Martes americana) may be i 8 impacted by trapping when ac- 5 ZF cess is high and avoid crossing large open areas, which could be 4 i impacted by road density (Chapin et al. 1998). Other car- t5 2 nivores that require large home ranges may also be affected by 1 road density. 0 4.43 0.434.63 0.850.84 0.84-1.14 1.141.55 Road Densky (KmlKmY) Management and research recommendation Figure 1. Cause of mortality of radio-collared wolves in Wisconsin 1979 to 1999, in Road density appears to be an relation to road density. important habitat factor for wolves and other carnivores; therefore public lands managed for these species should main- pack for over six years. Strict con- tain suitable habitats with low den- Discussion trol on access, and human activity sities of roads. Forested areas man- As human attitudes toward wolves limited to day light hours, can nul- aged for wolves should maintain improve and wolf populations con- lify the effect of road density. In overall road densities of 0.6 km/ tinue to increase, road densities Wisconsin, packs occur in the Crex km2. Core wolf habitat should be may become less of a factor in wolf Meadow Wildlife Area and managed at road densities of 0.45 habitat selection (Mech 1995). In Necedah National Wildlife Refuge km/km2. If productive packs exist the Great Lakes region, road den- that are classified as low probabil- in areas at much higher road den- sity provides a useful proxy for ity of pack occupation, but stricter sity, a reduction to lower densities human disturbance and risk of mor- access control by the management would not be necessary, but at- tality. In mountainous terrain agency reduces the effects of high tempts should be made to avoid in- where ungulate distribution is very road densities. Wolves do not have creasing road density or changing patchy, road density may be a less an aversion to roads, and readily traffic levels. useful index (Diane Boyd, personal travel on roadways if traffic levels On public forest lands, new log- communication), but in the gener- are low (Gehring 1995). Wolves ging roads should be closed or ally homogeneous landscape of the learn to avoid roads with high traf- obliterated after logging operations northern Great Lakes region it con- fic volumes, but readily use gated are completed. Where possible, tinues to be useful. Whether road roads (Thurber et al. 1994). Al- temporary, winter-only roads density serves as a good predictor though wolves seem to cross sec- should be used, because these roads of wolf habitat in northeastern U.S. ondary roads, they vary in willing- cause least damage and revert back remains to be seen. ness to cross busy highways (Frair to vegetated areas. Areas within Exceptions to the usefulness of 1999). Some dispersers do exten- 100 meters of den sites should be road density will continue to occur. sive crossings of highways (Kohn kept undeveloped, and logging Mech (1989) cites an example of ar- et al. 2000; Mech et a1 1995; Merrill roads and trails should stay more eas with road densities of 0.73 km/ and Mech 2000). than 100 meters from those sites. km2 supporting wolf packs in Min- Road density as a habitat factor The impact of road density is nesota but having large wilderness has applicability to other carnivores. not well known for other carni- reservoirs nearby. Merrill (2000) Roads affect movements and harvest vores; therefore research on habi- indicates that a military base in of black bear populations (Ursus tat use should include assessments Minnesota with a road density of americanus) (Brody and Pelton of road density as a habitat value. 1.42 km/km2 has supported a wolf 1989). (Lynx rufus) avoid The impacts of various types of

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Voi. 18 No. 4 2001 Endangered Species UPDATE 113 roads on carnivores should be studied and impacts from all-ter- rain vehicles (ATVs), snowmo- biles, and other off-road vehicles should also be investigated. Roads are important habitat vari- ables for carnivores that need to be more carefully researched.

Literature cited Berg, B. and S. Benson. 1998. Summary report 1997-1998 Wolf Survey Coop- erators. Minnesota Department of Natu- ral Resources. 4 p. (unpublished report). Brody, A.J. and M.R. Pelton. 1989. Ef- fects of roads on black bear movements in western North Carolina. Wildlife Society Bulletin 175-10. B~~~,M. and M. de Almeida, 1998. A re- Figure 2. Human causes of mortality of radio-collared wolves in Wisconsin 1979 view of wolfand coyote status and policy to 1999 in relation to road density. in Ontario. Ontario Ministry of Natural Resources. Pewborough Ontario. Chapin, T.G., D.J. Harrison and D.O. Katnik. 1998. Influences of landscape pattern on habitat use by American marten Lovallo, M.J. mdE.M. Anderson. 1996. Bob- in an industrial forest, Conservation Biol- movements and home range relative to lation levels. Journal of Wildlife Manage- ogy 12:1327-1337. roads in Wisconsin. Wildlife Society Bulle- ment 62: 1- 10. Corsi, F.,E. Dupre, and L. Boitani. 1999. A tin 24:71-76. Mladenoff, D.J, T.A. Sickley and A.P. large scale model of wolf distribution in Italy Massolo, A. and A. Meriggi. 1998. Factors Wydeven. 1999. Predicting gray wolf for conservation planning. Consewation Bi- affecting habitat occupancy by wolves in landscape recolonization: logistic regression ology 13: 150-159. northern Apennines (northern Italy): a model models vs. new field data. Ecological Ap- Frair, J,L. 1999. Crossing paths: gay wolves of habitat suitability. Ecography 21:97-107. plication 9:37-44. highways in the M~nesota-Wisconsin Mech, L.D. 1989. Wolf population survival in Shelley, D.P. and E.M. Anderson. 1995. Impact border region. University of Wisconsin, area of high road density. American Mid- of U. S. Highway 53 expansion on timber Stevens Point, WI MS. Thesis. 57 pp. land Naturalist 125387-389. wolves - Baseline Data. University of Wiscon- Fuller, T.K., W.E. Berg, G.L. Raddle, M.S. Mech, L.D. 1995. The challenge and oppor- sin, Stevens Point, WI, Final Report. 32 pp. Lenarz, G.B. Joselyn. 1992. A history and tunity of recovering wolf populations. Con- %el, RR. 1985. Relationshipbetween roadden- current estimate of wolf distribution and senation Biology 9:270-278. sities and wolf habitat suitability in Wisconsin. numbers in Minnesota. Wildlife Society Mech, L.D., S.H. Fritts, G.L. Radde, and W.J. AmericanMi~N~t113:40447. Bulletin 20:42-55. Paul. 1988. Wolf distribution and road den- Thompson, D.Q. 1952. Travel, range, and food Gehring, T.M, 1995, Winter wolf movements siQ in Minnesota. Wildlife Society Bulletin habits of timber wolves in Wisconsin. Jour- in northwestern Wisconsin and east-central I(i:@-87. nal of Mammalogy 33:429-442. Minnesota: a quantitative approach. uni- Mech, L.D., S.H. Fritts, and D. Wagner. 1995. Thurber, J.M., R.O. Peterson, T.D. Drummer, versity of Wisconsin, Stevens Point, WI Minnesota wolf dispersal to Wisconsin and and S.A. Thomasma. 1994. Gray wolf re- M.S. Thesis 132 pp. Michigan. American Midland Naturalist sponse to refuge boundaries and roads in Harrison, D. J. and T. G. Chapin. 1998. Ex- 133368-370. Alaska. Wildlife Society Bulletin 22:61-68. tent and comectivi~of habitat for wolves Menill, S.B. 2000. Roaddensities andgray wolf Unger, D.E. 1999. Timber wolf den and ren- in eastem ~~~hAmerica, Wildlife Society (Canis lupu~),habitat suitability: exception, dezvous site selection in northwestern Wis- Bulletin 26:767-775. CandianFieldNaturalist 114:312-313. consin and east-central Minnesota. Univer- Jensen, W,E, T,K, Fuller, W,L, Robinson, Menill, S.B. and L.D. Mech. 2000. Details of sity of W~SCO~S~,Stevens Point, WI. M.S. 1986. wolf (Canis lupus) distribution on the extensive movements by Minnesota wolves Thesis. 76 pp. OntarieMiChiganbordernearSaultSte.Marie. (Canis lupus). American Midland Natural- Wydeven, A.P., R.N. Schult~,and R.P. Thiel. Canadian Field-Naturalist 100:363-366. ist 144428-433. 1995. Gray wolf (Canis lupus) population Kohn, B,w,, J,L, Frair, D,E, Unger, T.M, Mladenoff. D.J., T.A. Sickley, R.G. Haight, and monitoring in Wisconsin 1979-1991. Pp. Gehng, D.P. Shelley, E.M. Anderson, and A.P. Wydeven. 1995. Aregiond landscape 147-156 in L. N. Carbyn, S. H. Fritts, and P.W. Keenlance. 2000. hpact of (he U.S. analysis and prediction of favorable gray D. R. Seip, eds. Ecology and conservation Highway 53 expansion project on wolves in wolf habitat in the Northern Great Lakes re- of wolves in a changing world. Canadian northwestem Wisconsin, Final Report for gion. Conservation Biology 9:279-294. Circumpolar Institute, University of , Wisconsin Department of Transportation, Mladenoff, D.J. and T.A. Sickley. 1998. AS- Edmonton, Canada. Wisconsin Department of Natural Re- sessing potential gray wolf restoration in the Wydeven, A.P. and J.E. Wiedenhoeft. 2000. sources. 49 pp + appendices. northeastern United States: a spatial predic- Gray wolf population 1999-2000. Wiscon- tion of favorable habitat and potential popu- sin Wildlife Surveys 10: 130-137.

114 Endangered Species UPDATE Val. 18 No. 4 2001 A Predator-Habitat Assessment for Felids in the Inland Atlantic Forest of Eastern Paraguay: A Preliminary Analysis

Gerald L. Zuercher Ph.D. Candidate, Kansas Cooperative Fish and Wildlife Research Unit, Kansas State University, 205 Leasure Hall, Manhattan, KS 66506; (785) 532-6336; (fax) (785) 532-7159; Research Associate, Sunset Zoological park; geraldza ksu.edu Philip S. Gipson Unit Leader, Kansas Cooperative Fish and Wildlife Research Unit, Kansas State University, 205 Leasure Hall, Manhattan, KS 66506; (785) 532-6070; (fax) (785) 532-7159; gipson Q ksti.edu Kim Hill Associate Professor, Department of Anthropology, University of , Albuquerque, NM 87131 ; (505) 277-1536; (fax) (505) 277-0874; kimhillQunm.edu

Abstract Jaguar (Panthera onca), puma (Puma concolor), and possibly six species of small cats (ocelot, Leopardus pardalis; margay, Leopardus wiedii; oncilla, Leopardus tigrinus; Geoffmy's cat, Oncifelis geoffroyi, pampas cat, Oncifelis colocolo, , Herpailurus yagouaroundi) co- exist within Mbaracayli Forest Nature Reserve in eastern Paraguay. At the landscape level, this 64,000-hectare island of Inland Atlantic Forest, surrounded by agricultural land, is a mosaic of forest habitats and interspersed . Habitats on the reserve include low, medium, and high forest, as well as dry and wet grasslands. The adaptive nature of most predators led us to predict that felids would occur uniformly across habitats. Tests of independence between species and habitats, howevel; suggest distinct associations between felids and habitats. Explanations for habitat afinities include integerence competition or simply following prey to their preferred habitats. Thus, while felids in the inland Atlantic forest may be habitat generalists across their entire range, they exhibit some habitat preferences within Mbaracayli reserve, possibly as an adaptation to interspecific competition and/or prey availability, Further research is needed to determine whether these patterns continue long-term or are an artifact of the timing of our current data collection efforts.

Reserva Natural del Bosque terized by extreme unpredictability have exceptional knowledge of the Mbaracayu in monthly patterns from year to Paraguayan forest because they The Mbaracayd reserve is the larg- year. The typical dry season lasts lived off wild resources until re- est tract of undisturbed forest in from May to September (Sanchez cently and most adults have spent eastern Paraguay covering approxi- 1973; FMB unpublished data). most of their lives in the forest (Hill mately 64,000 hectares of the quar- Seasons are associated with marked and Hurtado 1989). Further, the Ache ter million hectare upper Jejui wa- temperature fluctuations, with av- are allowed exclusive rights to hunt tershed. Only the combined reserve erage daily high-low temperatures and forage within the reserve. areas in Brazil and Argentina of 14 to 25 C in July and 22 to 34O around Yguazu Falls contains more C in January. The reserve contains Felid species within of the Alto Parana formation of the about 90% of all species classified Mbaracayu Reserve Atlantic Rainforest. Located at ap- as rare or endangered within Para- The diversity of mammalian carni- proximately 55O west and 24"south, guay (FMB 1992) and was chosen vores (Mammalia, ) exist- the reserve is mostly between 150 as the top priority conservation site ing within the bounds of the reserve to 300m in elevation and is drained in eastern Paraguay (Keel et al. is high with published lists of spe- to the west by the Paraguay River. 1993). The Mbaracayd reserve is cies ranging between 13 to 17 (Hill The area averages about 1800 mm located in the traditional homeland et al. 1997; FMB 1997; MNHNP of rainfall per year but is charac- of Ache hunter-gatherers. The Ache 1996). There is disagreement about

Vol. 18 No. 4 2001 Endangered Species UPDATE 115 the occurrence of some species, tors in much of their range. Pumas with the limited number of local with the following listed: three or are also large cats (males weighing names for small cats in the area four canids [bush ( as much as 103 kg); they are consid- (Table 2), led us to consider all venaticus), ered deer specialists, although they small cats as a group. (Chrysocyon brachyurus), crab-eat- feed on a diverse prey spectrum. ing fox (Cerdocyon thous), pampas Several species of small cats Major habitat types within fox (Pseudalopex gymnocercus)], potentially co-exist within Mbaracayu Reserve two procyonids [South American Mbaracay6 Reserve. The presence Until recently, categorizing the for- ( nasua), crab-eating of the ocelot, oncilla, jaguarundi, est mosaic of Mbaracay6 was dif- raccoon ( cancrivorus)], and Geoffroy's cat at this site is ficult. Sources for identifying in- four or five mustelids [ (Eira confirmed by voucher specimens at dividual plants and some plant barbara), lesser grison ( the Natural History Museum of communities have been available cuja) long-tailed ( Paraguay (MNHNP 1996). Mar- for reference for several years longicaudus), gays are considered residents (Hill (Lopez et al. 1987) but no consis- (Pteronura brasiliensis), hog-nosed et al. 1997) and pampas cat is ru- tent description of key habitat types (Conepatus chinga)], and, mored to exist in the cenado por- was available. The publication of five to eight felids [jaguar tion on the western edge of the re- "Plantas Comunes de Mbaracay6" (Panthera onca), puma (Puma serve (E. Esquivel, personal com- provides a scientifically based habi- concolor), ocelot (Leopardus munication). This discrepancy of tat classification system for studies pardalis), margay (L. wiedii), confirmed felid species, coupled within Mbaracayti (Marin et al. oncilla (L. tigrina), jaguarundi (Herpailurus yagouaroundi), Table 1. CITES listings for mammalian carnivores in Paraguay (MAG- Geoffroy's cat (Oncifelis geoffroyi), CITES 1999). pampas cat (0. colocolo)]. Many Carnivore Species Sciitific Name CITES Listing of these carnivore species, includ- 1 I I I ------ing all canids and felids, are listed by the Convention on International Trade in Endangered Species of bush dog Speothos venaticus Appendix I Wild Fauna and Flora (CITES), ei- maned wolf Chrysocyon brachyurus Appendix II ther on Appendix I or Appendix I1 (Table 1). We chose to focus on pampas fox Pseudalopex gymnocercus Appendix II felids as a first analysis of habitat associations by predators in Mbaracay6 reserve. The jaguar is found in many Appendix I habitats throughout its range (southwestern United States through northern Argentina). This is the largest native cat in the west- ern hemisphere with males weigh- jaguar Panthera onca Appendix I ing as much as 136 kg. Jaguars Puma concolor Appendix I1 are a top predator, and, in Mbaracayd Reserve, evidence of ocelot Leopardus pardalis Appendix I predation by a jaguar on a puma has been documented (K. Hill, personal I margay 1 Leopardus wiedii 1 Appendix l I observation). Jaguars are thought Appendix I to be dietary generalist and take ad- vantage of any prey available. Pu- mas currently exist over a larger area (northern Canada through southern Chile and Argentina) than pampas cat Oncifelis colocolo Appendix ll jaguars and function as top preda- I 1 I I

116 Endangered Species UPDATE Vol. 18 No. 4 2001 Table 2. Indigenous and local common names for possible small cat tree species > 10cm dbh is 39m2per species in ~eservaNatural del Bosque Mbaracayu. No Ache names ha in high forest (Keel 1987). exist to distinguish Margay, Geoffroy's cat, or Pampas cat. Jaguarete'i, and Tirika are each used for multiple cat species (Villalba Ground cover in high forest is gen- and Yanosky 2000, MAG-CITES 1999, FMB 1997). erally sparse including ferns, English Common Name heliconias (Heliconia psiattacorum; Musaceae), and bromeliads. Ap- ocelot Jaguarete'i, Jaguatirika proximately 30% of the Reserve area margay Jaguarete'i, Tirika is comprised of high forest. oncilla Kajamini Tirika Grasslands Geoffrey's cat Tirika, Tirika'i Grasslands are areas of open graminoid (i.e., Loudetiaflammida, pampas cat Osio Axonopus suflultus, Andropogon bicornis) meadow with patches of jaguarundi I Jaguarundi, Eira cerrado-like vegetation (i.e., palms, Butia paraguayensis; Palmae). They are usually wet during part of the year and are characterized by 1998). Habitat descriptions pro- and the tree height associated with poor drainage and a layer of black vided by the Ache were synthesized small bamboo understory ranges organic matter overlaying white into the categories outlined in from 15 to 25 m. Copaifera sandy soils. Several leguminous Marin et al. (1998). In our treat- langsdorfii or Hexachlamys edulis species (Mimosa spp., Desmodium ment of habitats within Mbaracayd can be the dominant woody tree spp., and Chamaecrista spp.) also we recognize three (low, medium, species. Other portions of medium characterize this habitat. Wet grass- and high) forest types, grasslands, forest are dominated by lianas. lands are dominated by several and cerrado. These areas are typically far from grasses (Setaria paucifolia and a permanent water source and Paspalum plicatum; Gramineae and Low forest ground cover is sparse. Although Rhynchospora glabosa; Located on dry land above stream riparian forests are distinguished by Cyperaceae), some of which can drainages, low forests are charac- the Ache as well as in several treat- reach two meters in height. These terized by trees <15 m high with a ments of the reserve, here it is in- areas are wet throughout the year typical diameter at breast height cluded in medium forest for the pur- with standing water up to half meter (dbh) <10 cm and none with a dbh pose of analyses. These areas are deep during some parts of the year. >25 cm. The understory is dominated in the downslope near streams and by bromeliads (Bromeliaceae) and rivers and are generally richer in Cerrado ferns (Blechnaceae, Cyatheaceae). fruit species. As with low forest, A large area of true cerrado habitat Major fruiting plants within low medium forest covers approxi- (-7,000 hectares) now exists within forest are Faramea porophylla and mately 30% of the Reserve area. the reserve. Cenado is a dry grass- Coussarea contracta within the landtforest interface dominated by Rubiaceae. Low forest constitutes High forest mixed-height grasses (Annona approximately 30% of the area of High forest, the most common plant coriacea, Dugetia furfuracea; the Reserve. community within Mbaracayd re- Annonaceae, and Eragrostis serve, is described in detail in Hill polytricha; Gramineae) and palms Medium forest et al. (1996). The canopy of this (Butia paraguayensis, Palmae). Be- Medium forest is characterized by forest is relatively tall (typically 12 cause this area was not a part of the variability. Some portions of me- to 20 m with some trees exceeding reserve when data collection was ini- dium forest are dominated by bam- 30 m) and trees commonly have tiated, however, it has been excluded boo, either the large Guadua epiphytic orchids (Orchidaceae) from surveys since its purchase. angustifolia bamboo or the small and philodendrons (Araceae). It is diameter Merostachys clausseni found at intermediate elevations Research question bamboo. There are few trees asso- above the water table on gently Given the information at hand, we ciated with the large bamboo stands sloping ground. The basal area of sought to determine if there are pat-

Vol. 18 No. 4 2001 Endangered Species UPDATE 117 terns of association between spe- ceiver. Data were then categorized quires further investigation. Jag- cific felids and major habitat types. according to species (jaguar, puma, uars appear to be the exclusive top We hypothesize that: felids in small cat) and habitat type prior to predator in Mbaracayd reserve; Reserva Natural del Bosque statistical analysis. In order to de- their diet overlaps and includes all Mbaracayd exhibit no detectable tect deviations from random habi- other cat species. habitat associations. Our rationale tat associations, a Chi-square test Pumas occur within all habitat for this are (1) predators are gener- for independence between species types as expected. The lack of a ally highly adaptive, and (2) the occurrence and habitat was per- pattern by pumas may reflect an habitats in Mbaracayd exist as a formed using SAS (SAS Institute overall high abundance of brocket mosaic with no distinct patches Inc., Cary, NC 27513). Data were deer, Mazama spp., within available for exclusive residence. pooled across survey years to en- Mbaracayd reserve (Hill unpub- sure a robust analysis. lished data). Data collection and analysis Small spotted cats occur within A stratified random sample of walk- Results and discussion low forest and habitats ing transects through the The Chi-square test resulted in a greater than expected and in high Mbaracayd reserve was employed significant deviation from random forest habitat less than expected. as the field method of game (x" 22.29, P = 0.0012) indicating The apparent preference for low censusing. Data for this analysis that some species were non-ran- forest and grasslands by small spot- come from four census periods: domly associated with habitats. ted cats may reflect an avoidance June 1994 to April 1995, July 1995 The relationship between species of jaguars, and other large preda- to June 1996, July 1996 to May occurrence within each habitat is tors, or a response to more abun- 1997, and July 1997 to May 1998. displayed in Figure 1. Jaguars oc- dant prey. Data for small Five Ache assistants, walking in cur within high forest habitat and small population densities parallel, and a data recorder, walk- greater than expected and in low are not available. ing 5 m directly behind the middle forest and grassland habitats less Thus, while felids in the inland researcher, censused the same 91 than expected. This apparent pref- Atlantic forest may be habitat gen- transects each year of the study. erence for high forest by jaguars eralists across their entire range, One individual walked directly on may be related to greater availabil- they exhibit some habitat prefer- the transect line while two assis- ity of prey species; an issue that re- ences within Mbaracayd reserve, tants were located on each side, separated by 25 and 50 m. Each transect began at a specified GPS location and proceeded along a specified compass bearing through------out the day for approximately 5 km. Transects passed through all habi- tat types and were only suspended when water exceeded half meter in depth. Transects were continued on I the far side of water bodies (swamp, streams, river, etc.) that had to be crossed. A GPS location and habi- tat description were noted for all types of animal encounters (for all animals at least 0.5 kg in size). The type of sighting was noted: (1) ani- mal was seen, heard, or found in a Erpoctrd Acturl wAch.l Expected A- EIpseted Aehlrl burrow, or (2) identified through mgb Fomt M* Pomt Lon Fomt Gmdradr fresh sign or feces. All locations 8J-r UPama ESmrUCats during the study were determined using a Trimble Pathfinder Pro Glo- Figure 1. Expected and actual counts of jaguars, pumas, and small cats within bal Positioning System (GPS) re- four habitat types in Mbaracayu reserve.

118 Endangered Species UPDATE Vol. 18 No. 4 2001 possibly as an adaptation to prey Paraguay. Pp 79-105 in J. Robinson and MAG-CITES (Ministerio de Agricultura y availability and interspecific com- E. Bennett, eds. Hunting for Ganaderia - Oficina CITES-Paraguay), sustainability in tropical forests. Colum- 1999. Especies Nativas de Paraguay. petition. Research is needed to test bia University Press, New York. MAG-CITES, Asunci6n, Paraguay our hypothesis that small cats ap- Hill, K., J. Padwe, C. Bejyvagi, A. Marin, G., B. Jiminez, M. Pena-Chocarro, pear to be associated with distinct Bepurangi, F. Jakugi, R. Tykuarangi, and and S. Knapp. 1998. Plantas Comunes habitats, in response to potential T. Tykuarangi. 1997. Impact of hunt- de Mbaracay6. The Natural History Mu- predation by jaguars and pumas. ing on large vertebrates in the seum, London. Mbaracayd Reserve, Paraguay. Conser- MNHNP (Museo Nacional de Historia vation Biology 11:1339-1353. Natural del Paraguay). 1996. Literature cited Keel, S. 1987. Informe de viaje a1 terrano Colecciones de Flora y Fauna del Museo FMB (Fundacidn MoisCs Bertoni). 1992. del Banco Mundial. Centro de Datos de Historia Natural del Paraguay. Reserva Natural Mbaracayd. Plan opera- para la Conservacion, Asunci6n, Para- Direccion de parques Nacionales y Vida tive. Fundacidn Mois6s Bertoni, guay. Silvestre, Ministerio de Agricultura y Asuncidn, Paraguay. Keel, S., A. Gentry, and L. Spinzi. 1993. Ganaderfa, Asunci6n, Paraguay. FMB (Fundaci6n MoisCs Bertoni) 1997. Using vegetation analysis to facilitate Sanchez, T.F. 1973. The climate of Para- Reserva Natural del Bosque Mbaracayd. the selection of conservation sites in guay. Pp 33-38 in J.R. Gorham, ed. Plan de manejo 1997-2001. Fundacidn eastern Paraguay. Conservation Biology Paraguay: ecological essays. Academy MoisCs Bertoni, Asuncidn, Paraguay. 7:66-75. of the Arts and Sciences of the Ameri- Hill, K. and A. Hurtado. 1989. Hunter- Lopez, J.A., E.L. Little, Jr., G.F. Ritz., J.S. cas, Miami. gatherers of the New World. American Rombold, W.J. Hahn, W.R. Sterrett, and Villalba, R. and A. Yanosky. 2000. Guia Scientist 77:436-443. M.J. McDonald. 1987. Arboles de Huellas y Seiiales: Fauna Paraguaya. Hill, K., J. Padwe, C. Bejyvagi, A. Comunes del Paraguay. Cuerpo de Paz, Fundacidn MoisCs Bertoni and U.S. Bepurangi, F. Jakugi, R. Tykuarangi, and Coleccidn e Intercambio de Agency for International Development, T. Tykuarangi. 2000. Sustainability of Informaci61-1, Asuncidn, Paraguay. Asuncidn, Paraguay. Ache hunting in the Mbaracayd Reserve,

The FLORIDA PANTHER ( concolor coryi) is a subspecies of mountain lion. However, this endangered cat resides in isolated areas of the Florida Everglades and Big Cypress Swamp. The males measure seven feet in length (including the tail) and weigh 100 to 160 pounds. Females are smaller at six feet and 55 to 100 pounds. The Florida panther is mostly crepuscular (active at dusk and dawn) and feeds primarily on white-tailed deer but also con- sumes wild hogs and raccoons. As habitat shrinks, the remaining population is experiencing a loss of genetic viability making it vulnerable to disease. By donating time or money to a nature conservation organization you can help save its habitat and help with relocation efforts and captive-breeding programs. 01999 by endangered species artist Rochelle Mason www. rmasonfinearts.corn (808) 985- 731I

Vol. 18 No. 4 2001 Endangered Species UPDATE 119 Raptor Conservation Status of the California Condor and Mortality Factors Affecting Recovery Kelly J. Sorenson Wildlife Restoration Coordinator and Assistant Director, Ventana Wilderness Society, P.O. 894, Carmel Valley, CA 93924; (831) 455-9514, (fax) (831) 455-2846; [email protected] L. Joseph Burnett Condor Field Coordinator, Ventana Wilderness Society, P.O. 894, Carmel Valley, CA 93924; http://w.ventanaws.org James R. Davis Executive Director, Ventana Wilderness Society, P.O. 894, Carmel Valley, CA 93924; http://www.ventanaws.org

Abstract The California Condor Recovery Program (Program) is moving forward after the release of 116 California condors (Gyrnnogyps californianus) to southern and central California and northern Arizona. As of February 1, 2001 a total of 161 condors were alive of which 46 were in the wild: 24 in California and 22 in Arizona. Due to a high number of fatalities thus far (431, the Program remains challenging. Discovering the relative importance of present-day mortality factors and controlling especially problematic ones is critical to recovery efforts. The effects of lead toxicity as a result of contamination from spent rifle and shot ammunition, and possibly other sources, is the most signijtcant threatpresently. As a result,jield release programs must focus a great deal of effort on ways to counter mortality until long-term solutions to this problem can be realized. This paper discussesfrom the perspective of Ventana Wldemess Society, the current status of the Calijor- nia condor; mortality factors that affect recovery, and management strategies to reduce fatalities.

Current status of recovery two geographically separate wild, rity and therefore successful breed- program self-sustaining populations each ing may occur in the near future. In In 1987, all remaining wild condors having 150 individuals, 15 breed- order to attain a self-sustaining popu- were brought into captivity to pre- ing pairs and a positive rate of lation, controllable mortality factors vent their extinction and rebuild the growth as well as a third popula- must be identified and reduced. population. By 1992, captive tion with 150 individuals in captiv- breeding at the Zoological Society ity in order to qualify for reclassi- Mortality factors affecting of San Diego and Los Angeles Zoo fication from endangered to threat- recovery increased the population to 52 ened. As of February 1,2001, 116 California condors are a long-lived and U.S. Fish and Wildlife Service different condors have been re- k-selected species with low repro- (FWS) began releases of young leased at three different geographic ductive rates (Temple and Wallace condors to southern California. The locations and despite heavy losses 1989). Wild condor populations Peregrine Fund began captive breed- the wild population maintains a cannot sustain themselves with an- ing in 1993 and releases to Arizona fairly evenly distributed age struc- nual mortality rates exceeding five in 1996. Ventana Wilderness Soci- ture (Table 1). A total of 27 con- to 7.5% for adults and 13 to 15% ety began releases in central Califor- dors have been removed to captiv- for juveniles (Verner 1978; nia in 1997 to aid the recovery of the ity for possible re-release and an- Meretsky et al. 2000). Up to the species. Captive breeding has other 43 birds have died or were early twentieth century, the condor brought the total population size to presumed dead. A total of 46 con- population plummeted due to losses 161 birds as of February 1, 2001 dors remain in the wild-24 in Cali- primarily caused by shooting. poi- (Jurek personal communication). fornia and 22 in Arizona. The old- soning, and museum collecting The California Condor Recov- est members of the reintroduced (FWS 1996). Throughout the re- ery Program Plan (1996) calls for population are now entering matu- maining 1900s the condor contin-

120 Endangered Species UPDATE Vol. 18 No. 4 2000 Table 1. California Condor releases, losses, and age structure natural causes is virtually unknown in wild (as of February 1, 2001). (a) "Total" includes one bird (FWS 1996). The high rate of deaths (b). . relocated to Arizona and therefore was released twice. to predation is likely a result of the re- Southern Central Arizona Total introduced birds not having parental California California I I I I guidance or protection in the wild. No, released 48 22 47 117(a) 1 Shooting No, removed to captivity or 12 7 8 27 possible re-release Shooting greatly affected the origi- nal wild population since scores of No. relocated 0 1(b) 0 1 I I I I condor deaths were attributed to No. died or presumed dead 26 0 17 43 this factor both done indiscrimi- nately and for the purposes of mu- seum collecting (Wilbur 1978). In I No. I-year-olds in wild I 0 1 0 I 5 1 5 1 the reintroduced population only two condors were shot and killed. I NO. 2-yeawids in wild 111511171 While this threat may always loom I No. 3-year-olds in wild 1°1314171 over the recovery of this species, it does not, by itself, appear to be a I No. 4-year-dds in wild 1 2 1 6 1 2 1 10 1 significant problem at present. I NO. 5-year-olds in wild I 0 I 0 I 4 1 4 1 Lead poisoning 1 No. 6-year-olds in wild 1 7 1 0 1 6 1 13 1 Lead exposure and acute poisoning I Total in wild 1 10 ( 14 122 146 I was first detected in the original wild population of California con- ued to decline though the reasons the remaining individuals regularly dors during the 1980s. Between were not entirely clear. perched on power poles, which led 1984 and 1986 three condor deaths Known causes of death in the to their return to captivity (Snyder were attributed to lead poisoning reintroduced population include: and Snyder 2000). Beginning with and one bird showed a positive ra- power line collisions/electrocu- the 1995 release cohort, a negative diograph for a bullet fragment in its tions, coyote (Canis latrans) and conditioning program was initiated digestive tract (Janssen et al. 1986). (Aquila chrysaetos) to train condors from perching on Other evidence of the lead threat predation, lead poisoning, acciden- power poles (Wallace personal within the range of the species in- tal drowning, shooting, ethylene communication). The results of this cludes studies on condors (Wiemeyer glycol poisoning, aspiration, can- training appear to be promising. et al. 1988), turkey vultures cer, and malnutrition (Jurek per- Prior to aversion training, 31 % (Cathartes aura) and ravens (Coruus sonal communication). Meretsky et (four of 13) of the condors released corm) (Wiemeyer et al. 1986b), and al. 2000 determined that recent an- died to collision and or electrocu- golden eagles (Pattee et al. 1990). nual mortality rates of the reintro- tion with power lines, while only Condors encounter lead bullet/ duced population are in excess of 2% (two of 103) died of the same pellet fragments in their food sup- what is critical to maintain a self- cause after the training was re- ply, although the exact pathways sustaining population. In the fol- quired for all releases. As a result, are not well understood. Currently, lowing section, we will discuss cur- this threat was greatly reduced. The lead rifle ammunition, in unrecov- rent mortality factors to shed light long-term effectiveness of this ered carcasses and/or gut piles fed on their impact on recovery efforts. training, however, is unknown. upon by the condor, may be the pri- mary source of contamination. A Electrocution/collision Predation single ingested lead fragment can Power line collisions and electro- Coyotes and golden eagles combined be lethal (Risebrough personal cutions represented a significant killed eight young or inex~riencedre- communication). Four confirmed threat to the reintroduced popula- introduced condors. Although ravens lead poisoning fatalities were docu- tion of condors during the first two and golden eagles are known threats to mented among all reintroduced years of release efforts. A total of condor eggs and nestlings respectively birds. To date, many of the rein- 4 birds were lost to this hazard and (Snyder 1986), adult mortality from troduced condors over the age of

Vol. 18 No. 4 2001 Endangered Specles UPDATE 121 one year experienced elevated lev- through public education, promo- soon be available (Oltrogge per- els of lead in their blood at one time tion of viable non-lead ammunition sonal communication). Until then, or another. Since 1997, 14 condors alternatives and enhanced food sub- education is an important step that were successfully treated for acute sidy efforts, (2) to improve moni- should be expanded. Currently, lead toxicity using a technique toring efforts utilizing aerial and clean food subsidy is provided at called chelation therapy. satellite tracking, and (3) to oper- all release sites. In addition, we Without this intensive interven- ate facilities in the field for post- support placing clean food at spe- tion effort, lead poisoning would release management. In the long cific locations near known nest sites easily account for the greatest num- run, measures taken to reduce the as they are discovered. ber of deaths. So important is the availability of lead in the condors' In response to increasing losses lead threat resulting from spent food supply represents the best to the reintroduced population and a ammunition in the condors' food sup- chance for eventual full recovery. growing number of unknown deaths ply that the California Condor Recov- Ventana Wilderness Society and disappearances, Ventana Wilder- ery Team is now recommending that supports public awareness efforts to ness Society initiated an intensive FWS reduce lead contamination openly discuss the problems of lead (weekly) aerial tracking program for within condor range (Recovery Team poisoning in wildlife, especially all condors in California beginning Meeting February 14, 2001). within the range of the California in fall of 2000 to augment the ongo- condor. Overall, the general pub- ing ground tracking effort. The au- Other causes lic knows little of this problem. The thors also support the use of satellite The original wild population expe- use of viable, non-toxic ammuni- telemetry on key individuals, espe- rienced some fatalities from drown- tion should also be encouraged cially those actively feeding on their ing, leg-hold traps, and in one case when possible. Non-toxic shotgun own to discover sources of lead con- a collision with a vertical pipe used ammunition is already commer- tamination and other threats. as a surveyor marker (Koford cially available and non-toxic rifle Currently, condors are released 1953). No evidence exists to sup- ammunition, called Ultimet, may both with and without post-release port any of these "other causes" playing a consistent role in the de- cline of the original wild popula- tion (Wilbur 1978). Accidental drowning (2), ethylene glycol poi- soning (I), cancer (I), aspiration (I), and malnutrition (1) are each recorded causes of death in the re- introduced population. Although these causes combined may affect current recovery to some extent, in- dividually they do not appear to be significant problems.

Unknown causes and disappearances Out of 116 different condors re- leased since the onset of the rein- troduction program, a total of seven deaths of unknown causes and an- other 10 disappearances (presumed dead) occurred. These results un- dermine the ability of the Program to combat limiting factors.

Management strategies The authors support strategies to (1) California Condor, stud book M80, free-flying over Ventana Wilderness, Cali- reduce the threat of lead to condors fornia. Photograph by Kelly Sorenson

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122 Endangered Species UPDATE Val. 18 No. 4 2001 management facilities in the field Literature cited Snyder, N.F.R. and Helen Snyder. 2000. The depending- on the release site. The California Condor Recovery Team Meet- California Condor: A saga of Natural His- authors support the use of post-re- ing. February 13-14, 2001. Sari Diego tory and Conservation. Academic Press. Wild Animal Park, Escondido, California. Temple, S.A., and M.P. Wallace. 1989. lease management at 'On- Janssen, D.L., J.E. Oosterhuis, J.L. Allen, Raptors in the Modern World. Pp 247- dor release sites. Ventana Wilder- M.P. Anderson. D.G. Kelts. and S.N. 251 in Meyburg, B.U. and R.D. Chan- ness Society uses a "double-door," Wiemeyer. 1986. Lead poisoning in cellor, eds. World Working Group Birds walk-in tra~attached to ~0st-l-e- free-ranging California Condors. J. of Prey, Pica Press. I Amer. Veter. Med. Assoc. U.S. Fish and Wildlife Service. 1996. lease management facilities, that 189: 17. California Condor Recovery Plan. 1996. managers to Koford, C.B. 1953. The California Con- Third Revision. Portland, Oregon. capture previously released con- dor. National Audubon Society Re- Verner, J. 1978. California condors: sta- dors. Routine evaluations of blood- source Report No. 41-154. tus of the recovery effort. Gen. Tech. Rep. PSW-28, U.S. Forest Service, lead levels can be achieved with Meretsk~,Vicky J.3 Noel F.R. Snyder, Steven R. Beissinger, David A. Washington, D.C. minimal stress to individual con- Clendenen, and James W, Wiley. 2000. Wiemeyer, S. N. Jurek, R. M, and J. R. dors and reduced staff effort. BY Demography-.- of the California Condor: Moore. 1986b. Environmental contami- increasing the level of monitoring Implications for Reestablishment. Con- nants in surrogates, foods, and feces of both in terms of tracking and cap- servation Biology. 11(1):957-967. California condors (Gymnogyps californianus). Environ. Monit. Assess. tures, field crews may be able to Pattee, O.H., Bloom, P.H., Scott, J.M., and M.R. Smith. 1990. Lead hazards within 6:91-111. further reduce fatalities. the range of the California Condor. Con- Wiemeyer, S.N., Scott, J.M., Anderson, dor 92,931-937. M.P. Bloom, P.H. and C.J. Stafford. Acknowledgements Snyder, N.F.R. 1986. California Condor 1988. Environmental contaminants in This manuscript was improved af- recovery program. PP 56-71 in S.E. California Condors. J. Wildl. Manag. Senner, C.M. White and J.R. Parrish, 52:238-247. ter review from Bruce eds. Raptor Reserch Report No. 5: Rap- Wilbur, S.R. 1978. The California Con- Robert Risebrough, Dr. Michael tor conservation in the next 50 years. dor, 1966-1976: a look at its past, present Wallace, Salvatore Lucido, Piper Raptor Research Foundation, Provo, and future. US gish and Wildlife Serivce Magallanes, and Carla Bundrick. Utah. North American Fauna 72: 1-1 36.

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Vol. 18 No. 4 2001 Endangered Species UPDATE 123 Raptors as Vermin: A History of Human Attitudes towards Pennsylvania's Birds of Prey

Keith L. Bildstein Hawk Mountain Sanctuary, 1700 Hawk Mountain Road, Kempton, Pennsylvania 19529; (610) 756-6000 x235

~bstract Many species of raptors (hawks, eagles, and falcons) were considered vermin in Pennsylvania well into mid-twentieth century. Indeed, as recently as the 1930s and 1940s, even eminent conservationists were calling for the elimination of so-called harmful birds of prey. Raptors were unprotected in Pennsylvania throughout the eighteenth and nineteenth centuries, and in I885 a 50-cent bounty was placed on all species of raptors. Although this particular bounty was re- pealed several years late~other bounties on diurnal raptors occurred sporadically until 1951. Bounties on several species of owls remained inforce until 1969. Raptorprotection, focusing on so-called beneficial species, first occurred in 1937. Bird-eating hawks, howevel; received only partial protection until 1969, and not all owls were protected until 1972, when the Migratory Bird Treaty Act of 1918 was amended to include birds of prey. At least part of the change in attitudes towards raptors can be attributed to activities at Hawk Mountain Sanctuary, the world'sfirst refuge for birds of prey, which was founded in Kempton, Pennsylvania, in 1934. Over the past two decades, populations of Pennsylvania's raptors have rebounded from shooting and pesticide- era lows of the early and mid-twentieth century. Recently, many hunters and bird watchers in the state have suggested that populations of raptors may once again be too high. As a result, Pennsylvania's raptor conservationists again face some of the same human attitudes theirprede- cessors faced more than a century ago.

Introduction were calling the "fool hawk law" United States and several foreign sig- Raptors and other vermin (i.e, harm- (Hornaday 1914). Raptors remained natories ratified an amended Migra- ful or objectionable animals) were unprotected in the state until 1937, tory Bird Treaty Act of 1918. Since unprotected in Pennsylvania through- when all species of diurnal birds of then, all species of diurnal birds of out the eighteenth and nineteenth prey, except for three accipiters prey and owls have been protected centuries. Persecution of raptors in (sharp-shinned hawk, Accipiter in Pennsylvania. Here I detail how the commonwealth increased sub- striatus; Cooper's hawk, A. cooperii; shifts in attitudes, both within and stantially in the latter half of the northern goshawk, A. gentilis) first outside of the conservation commu- 1800s, when an overwhelming ma- received protection (Kosak 1995). nity, contributed to this history. jority of rural residents considered Unfortunately, the new law was not raptors highly injurious. By 1885, particularly popular among Raptors as vermin animosity toward predatory birds had Pennsylvania's hunters and farmers, Raptors have had close associations intensified to the point that the state and scant enforcement within the with humans throughout history. legislature placed a 50-cent bounty state continued to plague so-called Many longstanding human-raptor as- on the heads of all diurnal birds of "protected" species well into the sociations are positive, including the prey, as well as on all owls. During 1960s. A bounty established on use of raptors in falconry, as national the next two years, 180,000 scalps northern goshawks in 1929 was lifted emblems and symbols of strength and were sent to the state capital in Har- in 1951, but it was not until 1969 that courage, and as flagship species for risburg, by which time increased Pennsylvania granted this species, broader conservation concerns. Un- populations of destructive rodents along with sharp-shinned and fortunately, raptors also have been and insects, together with fraudulent Cooper's hawks, full protection. scorned and feared, usually out of claims and a drain on the state trea- Great horned owls (Bubo ignorance. Because of this they have sury, induced the Pennsylvania leg- virginianus) remained unprotected been and continue to be heavily per- islature to repeal what by then many statewide until March 1972, when the secuted (e.g., Burnham 1990; Zalles

124 Endangered Species UPDATE Vol. 18 No. 4 2001 and Bildstein 2000). Indeed, to paraphrase Paul Errington (1946), "whatever else may be said of raptors and their predatory habits, they cer- tainly do draw attention." Systematic efforts to ex- terminate birds of prey can be traced to seventeenth century England (Gensbol1984). At- tempts at eradication, how- ever, escalated substantially I I in the 1800s, when the advent Figure 1. A series of three Works Progress Administration posters from the late 1930s of breech-loading guns in- commissioned by the Pennsylvania Game Commission in an attempt to help educate creased the popularity of hunters regarding the identification of 'good', and therefore protected species of rap- tors (e.g., Red-shouldered Hawks and Peregrine Falcons), versus 'bad' hawks (e.g., small-game hunting and Northern Goshawks). Note that the 'bad' goshawk is labeled "unprotected." placed hunters in direct com- petition with raptors (New- ton 1990). Polls taken at the Pearson 1897). The last half of the Renowned conservationist Will- time of passage of the Pennsylvania nineteenth and first half of the twen- iam T. Hornaday's world of animal Scalp Act of June 1885, a law that tieth centuries, was an era of 'good' protection also included both "good" established a 50-cent bounty on birds (i.e. rodent-eating) and 'bad' (i.e. bird- and "bad" hawks: " .. . 'chicken hawk of prey, suggested that the Act was eating) hawks (Fisher 1893) (Figure or hen hawk' are usually applied to supported by more than 90% of the 1). The idea that individual hawks the red-shouldered [Buteo lineatus] public in most areas of the state. Not and, in some instances entire species or red-tailed species. Neither of these surprisingly, contemporary conserva- of hawks, were "chickenhawks," and is really very destructive to poultry, tionists, including the president of the as such merited destruction, extended but both are very destructive to mice, New York Zoological Society, Will- well into the conservation community. rats and other pestiferous creatures.. . iamT. Hornaday, who referred to the The writings of conservationists Neither of them should be de- Act as the "fool hawk law" of the era reveal the animosity held stroyed-not even though they do (Hornaday 1914), and even Pennsyl- for some raptors. Consider, for ex- once in a great while, take a chicken vania state veterinarian and author of ample, this passage from John Muir's or wild bird," however " [tlhere are Diseases and enemies of poultry, The story of my boyhood and youth several species of birds that may at Leonard Pearson, considered the (1913): "When I went to the stable once be put under the sentence of Scalp Act unjust, uneconomic, and to feed the horses, I noticed a big death for their destructiveness of use- simply wrong-headed (Pearson white-breasted hawk [most likely a ful birds, without any extenuating cir- 1897). Although the law was re- red-tailed hawk, Buteojamaicensis], cumstances worth mentioning. Four scinded in 1887, the commonwealth on a tall oak in front of our chicken of these are Cooper's hawk, the sharp- reinstated bounties on raptors in house, evidently waiting for a shinned hawk, pigeon hawk [or mer- 1913, and maintained them on some chicken breakfast.. . I ran to the lin, Falco columbarius] and duck species until well into the 1960s house for a gun, and when I fired, he hawk [or peregrine falcon, F. (Kosak 1995). fell.. . then managed to stand erect. I peregrinus] " (Hornaday 1913). fired again to put him out of pain. He Hornaday's distinction appears to 'Good' versus 'bad' hawks flew off.. . but then died suddenly in have been both moralistic and utilitar- Most conservationists who opposed the air, and dropped like a stone." ian: "The ethics of men and animals are the Scalp Act took exception to the Although the event that Muir re- thoroughly comparative.. . Guilty ani- Act's all-encompassing nature- lated took place when he was a young mals, therefore, must be brought to jus- American kestrels (Falco span~erius) boy in 1850s Wisconsin, the founder tice" (Hornaday 1922). By 1931, were targeted along with northern of the Sierra Club expresses no re- Hornaday had dropped the Merlin from goshawks-rather than to the notion morse when recalling it in his auto- the list of "pest" birds, apparently be- that some hawks merited destruction biography more than half a century cause of its rarity, but retained the oth- (Hornaday 1913, 1914, 1931; later (Muir 19 13). ers, along with the

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Vol. 18 No. 4 2001 Endangered Species UPDATE 125 (Bubo virginianus), barred owl (Strix first half of the twentieth century. Un- varia), and eastern screech-owl (Otus fortunately, because many of the shoot- asio) (Hornaday 1931). ers either were unwilling or unable to The broader ornithological and separate the 'bad' hawks from "good," birdwatching communities, too, took all species of raptors remained at risk aim at certain raptors during the first (Broun 1949; Kosak 1995) (Figure 2). three decades of the twentieth cen- The impact of such shooting was little tury. The prolific and highly regarded studied, probably because much of the bird artist, Louis Agassiz Fuertes, conservation community condoned or writing in National Geographic, for even favored it. Even so, banding re- example, commented that "The coveries for Cooper's Hawks suggest whole genus Accipiter, consisting of that first year mortality due to shoot- the [northern] goshawk, Cooper's ing ranged from 28 to 47% in 1929 to hawk, and sharp-shinned hawk, are 1940, and from 12 t021% asrecentlyas savage, bloodthirsty, and cold- 1946 to 1957 (Henny and Wight 1972). hearted slaughterers, and are respon- sible in large measure for the anath- The goshawk invasion of ema that is then portion of all hawks" the late twenties Figure 2. The results of raptor shoot- ing in Pennsylvania in the late 19509, (Fuertes 1920). And famed As hated as resident accipiters were, when Sharp-shinned Hawks (top) re- birdwatcher and president of the Con- migrants from the north were de- mained unprotected, while Ospreys necticut Audubon Society, Mabel spised even more (Gerstell 1937). (Pandion haliaetus) (bottom), were Osgood Wright, suggested helping Northern Goshawks, in particular, protected (photographs courtesy Hawk Mountain Sanctuary archives). songbirds by "shooting some of their were singled out in this regard. As enemies" including several species of ornithologists Edward H. Forbush re- hawks (Wright 1936). Similarly, marked in his Birds of Massachusetts and 1 May. Not surprisingly, the Pennsylvania's official "state orni- "A great flight of goshawks into the bounty substantially increased raptor thologist," and later president of the United States in fall or winter is fol- shooting at the site, as well as Wilson Ornithological Society, lowed invariably by a scarcity of throughout Pennsylvania. Second, George Miksch Sutton, wrote in the Ruffed Grouse [Bonasa urnbellus]" Sutton's articles alerted the ornitho- Introduction to the birds of Pennsyl- (Forbush 1929). Thus, when rural in- logical and conservation cornrnuni- vania that: " [tlhe sharp-shin is the en- habitants of Drehersville, at the base ties of both raptor migration and emy of all small birds.. . [and that it] of the Kittatinny Ridge in the central shooting at the site, which the locals and [the] Cooper's hawk, both bird Appalachian Mountains of eastern called Hawk Mountain. killers, are fairly common and are Pennsylvania, reported an "invasion" Earl Poole, then assistant cura- rated as our most objectionable birds of northern goshawks during the win- tor at the Reading Public Museum in of prey. They are not protected in ters of 1926-1927 and 1927- 1928 to nearby Reading, Pennsylvania, began Pennsylvania." (Sutton 1928a). the Pennsylvania Game Commission, visiting Hawk Mountain in 1929. His Even Boy Scouts were instructed the commission quickly dispatched description of a broad-winged hawk in the whys and wherefores of "good" state ornithologist George Miksch (B. platypterus) flight of 2,000 birds and "bad" hawks. George E. Hix, a Sutton to investigate. on 22 September 1932, represents the Brooklyn, New York, scoutmaster and Sutton published his initial find- first detailed account of raptor migra- Associate of the American Ornitholo- ings in the Wilson Bulletin in 1928 tion at the site (Poole 1934). Shortly gists' Union wrote in his Birds of prey (Sutton 1928b). His report, together thereafter, conservationists Henry H. for Boy Scouts " .. . [that] the benefi- with a second paper published three Collins, Jr., and Richard Pough vis- cial hawks are the larger, slower spe- years later (Sutton 1931), initiated a ited the ridge, confirming Sutton and cies, [and that] the smaller swifter series of events that eventually re- Poole's earlier accounts, and photo- hawks are the ones which are destruc- sulted in the creation of the world's graphing the slaughter that was then tive to wildlife. . . [and that these include] first Sanctuary for birds of prey. underway (Pough 1932; Collins the goshawk, Cooper's, sharpshinned, First, by 1929, the Pennsylvania 1933). Pough showed slides of his duck and pigeon hawks.. ." (Hix 1933). Game Commission was offering a photographs to a joint meeting of the Small wonder then that bird-eat- new five-dollar bounty on northern Hawk and Owl, Linnean, and Na- ing birds were heavily persecuted in the goshawks shot between 1 November tional Association of Audubon soci-

126 Endangered Species UPDATE Vol. 18 No. 4 2001 Thirty-five years of conservation and likely to be as ineffective as remov- education efforts at the Sanctuary, in ing a single Gray Squirrel (Sciurus part, resulted in statewide, year- carolinensis), others appear deter- round protection for all diurnal birds mined to "do something themselves" of prey, including the three "bird-kill- about the situation, including, a few ing" accipiters, in 1969 (Senner have suggested, shooting the hawk 1984). It was not, however, until the (Bildstein, personal observation). Migratory Bird Treaty Act of 1918 Research conducted at Hawk Figure 3. The view from Hawk Moun- was amended in March of 1972, that Mountain Sanctuary and elsewhere tain Sanctuary, the world'sfirst refuge all raptors, including great horned suggests that recent shifts in the rni- for birds of prey, in the 1940s. (Photo: and snowy owls, were protected in gration behavior of eastern popula- Hawk Mountain Sanctuary archives.) Pennsylvania (Kosak 1995). tions of sharp-shinned hawks may be related to an increasing tendency to eties meeting in New York City in A modern era? pause at bird feeders along migration autumn 1933. Rosalie Edge, the Although all raptor species remain routes in the northeastern United head of the Emergency Conservation officially protected in Pennsylvania, States (Viverette et al. 1996; Duncan Committee, was in the audience the and have for almost thirty years, their 1996). Furthermore, data collected evening Pough spoke. status remains controversial. As re- by participants in the Cornell Univer- The following June, Mrs. Edge, ac- cently as 1999, the Pennsylvania sity Laboratory of Ornithology's companied by Richard Pough and her Game Commission held hearings on Project Feederwatch indicate that son Peter, visited Hawk Mountain. a proposal from the President of the sharp-shinned hawks take more birds Shortly thereafter, Rosalie Edge leased Commission (game commissioners at birdfeeders than do feral cats the 565 acres that was to become Hawk are appointed by the Governor) re- (Dunn and Tassaglia 1994). Whether Mountain Sanctuary (Bildstein and garding "experimentally" controlling or not increased numbers of accipi- Compton 2000). Today, the Hawk populations of Red-tailed Hawks and ters at bird feeders are impacting re- Mountain Sanctuary Association, the Great Horned Owls on several wild- gional populations of songbirds and largest and oldest member-based rap- life management areas in an attempt other species feeding at these sites re- tor conservation organization in the to increase the survivorship of Ring- mains unclear, although evidence world, manages the Sanctuary, which necked Pheasants (Phasianus from England suggests that this may has now grown to more than 2,300 colchicus) (Riegner 1999). Although not be occurring (Newton et al. acres. Once a shooting gallery where a public hearing revealed widespread 1997). On the other hand, studies of thousands of raptors were killed each opposition to the proposal-that was American kestrels that overwinter in autumn, Hawk Mountain's Appala- later withdrawn-letters to the edi- farmlands surrounding Hawk Moun- chian lookouts now serve as both learn- tors of local newspapers also sug- tain suggest that recently increased ing center and biological field station gested a degree of public support for populations of Cooper's and, possi- for more than 80,000 visitors annually the idea (e.g., Riegel 1999). bly, sharp-shinned hawks, both of (Zalles and Bildstein 2000) (Figure 3). Indeed, many birdwatchers, es- which prey on kestrels, are affecting Unfortunately, the Pennsylvania pecially those maintaining backyard regional populations of this small fal- bounty on goshawks (reduced to two birdfeeders, continue to call Hawk con (Ardia and Bildstein 1997;Ardia dollars in 1937) remained in place Mountain Sanctuary to express out- et al. unpublished data). until 1951, when the state legislature rage at the seemingly persistent approved, and Governor G. H. Earle predatory activities of sharp-shinned Conclusion signed a law protecting all raptors, and Cooper's hawks at their bird feed- Raptor populations have increased excepting the three species of "bird- ers. Both species of accipiters ap- substantially in the last two decades killing" accipiters and two species of pear to be increasingly willing to en- of the twentieth century (Bednarz et owls, the great horned owl and the ter human-dominated landscapes; al. 1990; Bildstein 1998), quite pos- snowy owl (Nyctea scandiaca) most likely in response to reduced hu- sibly to levels similar to those of the (Kosak 1995). man-caused mortality there. Al- latter nineteenth and early twentieth The founding of Hawk Mountain though most callers seem resigned to centuries. In Pennsylvania and, in- Sanctuary highlighted raptor conser- this situation, particularly once they deed throughout most of North vation both within and outside of have been informed that removing a America, birds of prey are no longer Pennsylvania as never before. single accipiter from a backyard is the endangered boutique predators

Vol. 18 No. 4 2001 Endangered Spedes UPDATE 127 (i.e. predators whose populations are mal Behaviour 53: 1305-1311. red-tailed and Cooper's hawks. 4.229- 249 so low that they do not substantially Bednarz, J.C., D. Klem Jr., L.J. Goodrich, and in Population ecology of migratory birds. S.E. Senner. 1990. Migration counts of rap- Wildlife Research Report 2. U. S. Fish and influence the behavior and ecology tors at Hawk Mountain, Pennsylvania, as Wildlife Service, Washington, D. C. of their prey species) that they were indicators of population trends, 1934-1986. Hix, G.E. 1933. The birds of prey for Boy as recently as the late 1970s Auk 107:96-109. Scouts. Privately published, Brooklyn, NY. (Bildstein 1998). Now that raptors Bildstein, K.L. 1998. Long-term counts of Homaday, W.T. 1913. Our vanishing wild life. York Zoological York, NY. once again are fairly common and migrating raptors: a role for volunteers in New Society, New wildlife research. Journal of Wildlife Man- Hornaday, W.T. 1914. Wild life conservation fully functional components of natu- agement 62:435-445. in theory and practice. Yale University Press, ral and human-dominated land- Bildstein, K.L. and R.A. Compton. 2000. New Haven, CT. scapes, raptor conservationists are Mountaintop science: the history of conser- Homaday, W.T. 1922. The minds and mrs facing many of the same manage- vation omithology at Hawk Mountain Sanc- of wild animals. Scribner, New Yo& NY tuary. Pp. 153-181 in W.E. Davis and J.A. Hornaday, W.T. 1931. Thxty years war for ment concerns and human attitudes Jackson, eds. Contributions to the History wild life. C. Scribner's Sons, NewYork, NY. their predecessors faced more than of North American Ornithology, Volume II Kosak, J. 1995. The PennsylvaniaGameCom- one hundred years ago. Memoirs of the Nuttall Ornithological Club, mission 1895-1995. Pennsylvania Game As a result, keeping common No. 13. Nuttall Ornithological Club, Cam- Commission, Harrisburg, PA. raptors common at the beginning of bridge, MA. Muir, J. 1913. The story of my boyhood and Broun, M. 1949. Hawks aloft. Cornwall Press, youth. Houghton Mifflin, Boston, MA. the twenty-first century may prove to Cornwall, NY. Newton, I. 1990. Human impacts on raptors. be as much of a challenge for today's Burnham, W.A. 1990. Raptors and people. Pages 190-207 in I. Newton, ed. Birds of raptor conservationists (Garrott et Pages 170-189 in I. Newton, ed. Birds of Prey Facts on File, New York, NY. a1.1993) as it was for their predeces- Prey, Facts on File, New York, NY. Newton, I., L. Dale, P. Rothery. 1997. Appar- Collins, H.H., Jr. 1933. Hawk slaughter at ent lack of impact of Sparrowhawks on the sors at the beginning of the 20" Cen- Drehersville. Hawk and Owl Society Bul- breeding densities of some woodland song- tury. Experience at Hawk Mountain letin 3:lO-18. birds. Bird Study 44129-135. Sanctuary suggests that focused con- Duncan, C.D. 1996. Changes in the winter Pearson, L. 1897. Diseases and enemies of servation education that extends from abundance of sharp-shinned hawks in New poultry. Clarence M. Busch, Hanisburg, PA. primary schools through the general England. Journal of Field Ornithology Poole, E.L. 1934. The hawk migration along 67:254-262. the Kittatinny Ridge in Pennsylvania. Auk public, longterm population monitor- Dunn, E.H, and D.L. Tassaglia. 1994. Preda- 51:17-20. ing, and, above all, opportunities for tion of birds at feeders in winter. Journal of Pough, R.H. 1932. Wholesale killing of hawks viewing large numbers of these nor- Field Ornithology 658-16. in Pennsylvania. Bird-Lore 34429-430, mally secretive birds at migration Errington, P.L. 1946. Predation and vertebrate Riegel, E. 1999. No sympathy for killer hawks. populations. Quarterly Review of Biology Reading Eagle (newspaper). 11 March 1999. hawkwatches, are practical and effec- 21: 144-177. Riegner, C. 1999. Pheasant problems. Penn- tive ways to build local and regional Fisher, A.K. 1893. Hawks and owls of the sylvania Game 70(11): 16-18. support for our charismatic birds of United States in relation to agriculture. Senner, S.E. 1984. The model hawk law - 1934 prey. With this in mind, the Sanctu- U.S.D.A. Division of Ornithology and Mam- to 1972. Hawk Mountain News 6229-36. ary continues to work with local, re- malogy, Bulletin Number 3. U.S.D.A., Sutton, G.M. 1928a. An introduction to the Washington, D. C. birds of Pennsylvania. McFarland, Harris- gional, and national conservationists Forbush, E.H., ed. 1929. Birds of Massachu- burg, PA. to foster migration watchsites else- setts and other New England states, Vol. 11. Sutton, G.M. 1928b. Notes on a collection of where in the world so as to protect Massachusetts Department of Agriculture, hawks from Schuylkill County, Pennsylva- long-distance migratory raptors Boston, MA. nia. Wilson Bulletin 40: 84-95. Fuertes, L.A. 1920. American birds of prey - Sutton, G.M. 1931. The status of the Gos- throughout their journeys. a review of their value. National Geographic hawk in Pennsylvania. Wilson Bulletin 38:460-467. 43:108-113. Acknowledgments Ganott, R.A., P.J.White, C.A.V.White. 1993. Viverette, C.B., S. Struve, L.J. Goodrich, and I thank Kyle McCarty and Martin Smith Overabundance: an issue for conservation K.L. Bildstein. 19%. Decreases in migrat- biologists? Conservation Biology 7:946- ing sharpshinned hawks (Accipiterstriatus) for reviewing earlier versions of this 949. at traditional raptor-migration watch sites in manuscript, and Kev Biffna for scanning Gensbol, B. 1984. Birds of prey of Britain, eastern North America. Auk 113:32-40. the figures. This is Hawk Mountain Sanc- Europe, North Africa and the Middle East. Wright, M.O. 1936. Birdcraft, ninth edition. tuary contribution number 68. Collins, London, England. Macmillan, London, England. Gerstell, R. 1937. The Pennsylvania bounty Zalles, J. and K.L. Bildstein. 2000. Raptor system. Research Bulletin 1. The Pennsyl- watch: a global directory of raptor migra- Literature cited vania Game Commission, Harrisburg, PA. tion sites. Hawk Mountain Sanctuary Asso- Ardia, D.R. and KL. Bildstein. 1997. Sex-re- Henny, C.J. and H.M. Wight. 1972. Popula- ciation, Kempton, PA; and BirdLife Inter- lated differences in habitat selection in winter- tion ecology and environmental pollution: national, Cambridge, England. ing American Kestrels, Falco sparverius. Ani-

128 Endangered Species UPDATE Vol. 18 No. 4 2001 Felid Conservation Status and Conservation of Endangered Cats Along the U.S.-Mexico Border Melissa Grigione Deptartment of Environmental Science and Policy, College of Arts & Sciences, University of South Florida, 4202 East Fowler Ave., Tampa, Florida 33620; mmgrigione0 iname.com Arturo Caso Proyectos Sobre 10s Felinos Neotropicales de Mexico, Bosques 144-B, Lomas del Chairel, C.P. 89360 Tampico, Tam. Mexico; acaso20terra.com.mx Rurik List Institute de Ecologia, UNAM. 38rCircuito Exterior Ciudad Universitaria, Anexo al Jardin Botanico, Coyoacan, 04510 Mexico; rurik0toluca.podernet.com.mx Carlos Lopez-Gonzalez Department of Conservation Biology, Denver Zoological Foundation, 2300 Steel Street, City Park, Denver, Colorado 80205-4899: cats4rnexQaol.com

Abstract This paper reviews the research and conservation projects associated with the Bordercats Work- ing Group, a group of scientists and advocates concerned with the status of neotropical cats in northern Mexico and the American Southwest. For ocelots (Leopardus pardalis), jaguarundis (Herpailurus yaguarondi) and jaguars (Panthera onca), this region is the northernmost limit of their geographical distribution. We review the results of ourfield surveys for bordercats in Arizona, , Coahuila, New Mexico, Nuevo Leon, Sonora, Tamaulipas, and ; describe a GIs-based habitat mapping project for bordercats; and discuss what consewation- related activities are critical for the long-term survival of the cats in the border regions.

Introduction ment, and ranching practices that ternational laws, such as the There are three cat species whose are intolerant of predators. In ad- President's National Drug Control northernmost range coincides dition, the border region is a major Strategy, the National Defense Au- with the border region of the thoroughfare for illegal aliens. The thorization Act, and U.S.-Mexican United States and Mexico. These United States Border Patrol at- immigration policies, which often are North America's neotropical tempts to prevent such crossings by conflict with Federal environmen- cat species: the ocelot (Leopardus burning or mowing border vegeta- tal laws like the ESA and the Na- pardalis), jaguar (Panthera tion, constructing fences, and put- tional Environmental Policy Act onca), and jaguarundi ting up large floodlights across the (NEPA). In addition, Texas and (Herpailurus yaguarondi). Un- border, which for nocturnal cat spe- states in northern Mexico consist like the mountain lion (Felis cies are less than ideal. Recovery primarily of private land, making concolor) and bobcat (Felis for neotropical cat populations in a large refuge for the cats particu- rufus), these species are endan- the border region requires not only larly difficult. Despite this, out- gered and protected by both the an understanding of the spatial re- standing commitment to cat conser- U.S. Endangered Species Act quirements and habitat needs of the vation by the U.S. National Wild- (ESA) and by Mexican law cats but an understanding of how life Refuge system, particularly the (NOM-059- 1994), which prohib- the cats can best fit into a complex Lower Rio Grande National Wild- its hunting of the three species. human matrix consisting of farm- life Refuge (LRGNWR) and La- The border region presents an ing and ranching communities, guna Atascosa National Wildlife array of threats to neotropical cats, refugees, federal agencies, all con- Refuge, both in Texas, has elevated such as highway and international verging at the international border. public awareness about the cats and bridge construction, land develop- There are a suite of national and in- the challenges they face.

Vol. 18 No. 4 2001 Endangered Species UPDATE 129 Addressing conservation needs: The Bordercats Working Group The Bordercats Working Group was founded in 1998 to promote recov- ery and conservation for neotropical cats in the border region of the U.S. and Mexico. The group was created because of the great amount of biodiversity in the border region, evi- dence of few to no neotropical cats in this region in recent times, a gap in knowledge about the detailed dis- tribution of the species in this region, and an existing legal framework in place to protect all three species. Figure 1. Historical distribution of jaguarundls, laguars, and dots The Bordercats Working Group (BWG), now a member of the World Chihuahua, TexaslCoahuila, and consisted of cattle association offi- Conservation Union's (IUCN) Cat Tamaulipas/Nuevo Leon. cials, ranchers, and predator hunters. Specialist Group, has several objec- Whenever possible, physical evi- tives: to conduct research on all three Arizona/New Mexico dence (such as skins, skulls, and pho- species in the border region; to de- The Chiracahua and Peloncillo tographs) of recently killed cats was marcate important habitats for the Mountains were surveyed for cats for collected (Lopez-Gonzalez and cats in the border region; to educate four months during the Spring and Brown in preparation). local communities and organizations Fall of 1999. Field biologists used a There are approximately three ex- about the cats; and to bring individu- combination of techniques to detect tant metapopulations of jaguars in the als, groups, and government agencies cat presence: scat surveys, track sur- State of Sonora: Sierra Bacatete, together to share in a common mis- veys, remotely-triggered camera sur- Sahuaripa-Huasabas, and Quiriego- sion. This paper will focus on the veys, and formal interviews with bi- Sinaloa (Lopez-Gonzalez and Brown first two objectives: the distribution ologists and local naturalists. Dur- in preparation). Preliminary evidence of borderland cat populations, and the ing this survey, no physical evidence suggests that this area supports approxi- identification of important cat habi- of neotropical cats was obtained. mately 1.3 to 1.5 jaguars per 100 km2. tats in this region. There were, however, approximately Using a home range estimator (Adap- 12 interviews held with knowledge- tive Kernel model), a minimum of Distribution of bordercat able locals, including biologists, who 6,000 krn2would be needed to conserve populations attested to the presence of jaguarun- viable metapopulations in this region The historic distribution (Figure I) dis. More surveys would have to be (Brown and Lopez-Gonzalez 2000). for all three cat species in the border conducted to verify these reports. This initial research has found that at region is located in the U.S. South- least 140 jaguars have been killed in west (Arizona, New Mexico, and SonoraKhihuahua Sonora since 1900; 45 have been killed Texas) and adjacent Mexican States Approximately 215 km south of the since 1990 (Lopez-Gonzalez and (Chihuahua, Coahuila, Nuevo Leon, Arizona/New Mexico border, in the Brown unpublished report). A mini- Sonora, and Tamaulipas). At present, Mexican State of Sonora, lie impor- mum of 14 females, three lactating fe- there are approximately 100 ocelots tant areas for both jaguars and oce- males, and five kittens have been killed in the United States and no known lots; jaguarundis may occur just south since 1989. In 1999 to 2000, sixteen populations of jaguars or jaguarun- of this area. Surveys for all three spe- jaguars were killed in the northern dis. From 1982 to 1996,12 out of 17 cies, which incorporate formal inter- range of this distribution (Lopez- radio-collared ocelots in south Texas views and remotely-triggered cam- Gonzalez and Brown 2000). All of were killed by automobiles (Tewes eras, have been ongoing since June these animals were killed in response et al. 1997). BWG is involved with 1998. Formal interviews have oc- to livestock losses and none of the skins neotropical cat surveys in four re- curred in 50 municipalities in Sonora were sold internationally (Lopez- gions: Arizona/New Mexico, Sonora, and 12 in Chihuahua. Interviewees Gonzalez and Brown in preparation).

130 Endangered Species UPDATE Vol. 18 No. 4 2001 The closest known population numerous jaguarundi sightings from Identification of key bordercat of ocelots to ArizonaJNew Mexico park visitors but currently have no habitats is also in this area. From July 1998 physical evidence in support of this In order to identify what borderland to October 1999, 33 records of species inhabiting the park. habitats are most important to hunted ocelots were collected in neotropical cats, BWG initiated a this region; approximately 79% of TamaulipasNuevo Leon Geographical Information System these ocelots were killed after 1990 BWG has ongoing research in both (GIs) habitat mapping project in (Lopez-Gonzalez 2000). Although Tamaulipas and Nuevo Leon, just 1999. BWG collected sighting data three ocelots have been taken in south of the Texas border. Survey for each species, spanning the last Sonora within 40 km from the Ari- efforts have focused in areas, within 100 years. These sightings were plot- zona border, all were male. Due to close proximity to the border, which ted on a map containing the follow- an average dispersal distance of ap- have intact cat habitat and reports of ing information: vegetation, city and proximately 20 km, it is unlikely ocelots or jaguarundis (Figure 3). county boundaries, roads, rivers and that female ocelots would be found Once suitable areas are identified, a streams, and elevation. Mapping par- this close to the Arizona border. combination of track and scat sur- ticipants, each familiar with a particu- veys, remotely-triggered cameras, lar region or conservation unit (CU), Texas/Coahuila predator calls, and box traps baited were asked important questions about In March 1999, BWG surveyed the with chickens are used to detect cat each species sighting map. The ques- Rio Grande River near Big Bend presence. We are currently investi- tions pertained to both the species and National Park, in Texas and the Mexi- gating two recent reports of jaguarun- their habitats, major threats to each, can State of Coahuila for neotropical dis in close proximity to Monterrey, the development of conservation cor- cats. Approximately 100 km of river Nuevo Leon, approximately 200 km ridors between CU's, and the effec- and associated tributaries were sur- from the international border (Area tiveness of the land-tenure system in veyed during this trip. Transects #1, Figure 3). We have no evidence place within each CU. The results of walked along each sandy substrate of ocelots in this area; however, our this large mapping project will be adjacent to the river were scanned for surveys are not yet complete. We have synthesized and a final map will be cat tracks and scat. When tracks were surveyed in areas north of Monterrey produced. The final map will be found, they were measured and pho- and have not detected tographed, and plaster castes of each track set were made. A Global Posi- tioning System (GPS) unit was used to measure the location of each track set. In addition, data on substrate, time of day, and general habitat char- acteristics were recorded for each track set. Scent-stations were set up prior to sunset by sifting sand over one m2 areas, placed 10 meters from one another. Hawbawkers Cat Lure was spread over a stone and placed in the center of each one m2sampling unit. Each station was checked for cat tracks the following morning. During this survey, we observed several cat tracks, including nine mountain lion track sets (seven adults and two cubs), six bobcat track sets, and what appeared to be one jagua- rundi track set, all found at the mouth of the western end of the Boquillas Canyon (Figure 2). Big Bend Na- tional Park employees have collected habitat use. Figure 2. Details of bordercat sightings

Vol. 18 No. 4 2001 Endangered Species UPDATE 131 Foreman, Patricia Manzano-Fischer, Craig Miller, and Scott Royder dur- ing the Big Bend surveys was indispensible. John Morrison (World Wildlife Fund-U.S.A.), Eric

Kay sites Sanderson (Wildlife Conservation ** chlrryue rur re LyunoAtwora~W Society), Robert Thomas (Univer- 9. Lbr Elwr.Vh y LoStbmeS OW* sity of California-Davis), and Diana d L.mRlo OrncV#y NW 9 Irn(rtU0~Rnch , Lawhorn (The Nature Conservancy) ,* aa* Dom~Ranch are thanked for their involvement in the GIs Habitat Mapping Project. We wish to thank John Morrison and Robert Thomas for producing all maps contained in this manuscript.

Literature cited

0 Brown, D.E., and C.A. Lopez-Gonzalez. s 2000. Notes on the occurrences of jag- uars (Panthera onca) in Arizona and New Mexico. Southwestern Naturalist 45:537-542. Figure 3. Survey sites for neotropical cats in NE Mexico Caso, A. 1994. Home range and habitat use of three neotropical carnivores in North- east Mexico. M.S. Thesis, Texas A&M completed in Fall 2001 and will serve the status and distribution of bor- University - Kingsville. Lopez-Gonzalez, C.A. and D.E. Brown (in as a conservation blueprint for future derland cats. preparation). Distribution and current sta- priority research and habitat protec- tus of jaguars (Panthera onca) in North- tion projects related to neotropical Acknowledgements western Mexico. cats in the border region. We wish to thank all members of the Lopez-Gonzalez, C.A. 2000. Sonora ocelots: can they make it to Arizona? Journal of Bordercats Working Group for their the Arizona-Nevada Academy of Science: Conclusions ongoing participation, expertise, and Forty fourth annual meeting. The Univer- Obtaining valuable information commitment to cat conservation sity of Arizona-Tucson. about the distribution of neotropical along the U.S./Mexico border. We Lopez-Gonzalez, C.A., and D.E. Brown. cats and what habitats are most im- thank Defenders of Wildlife, Round 2000. Jaguars (Panthera onca) as conser- vation tools for Northwestern Mexico. portant to their conservation is a River Conservation Studies and the The Wildlife Society, Western SectionAb- first step toward promoting their Walt Disney Company Foundation stracts. Riverside Convention Center: Riv- long term persistence in the border for their financial assistance, in ad- erside, California. region. This research, however, dition to in-kind support from mem- Lopez-Gonzalez, C.A. and D.E. Brown (unpublished report to Wildlife Conser- needs to be married with effective bers of the Bordercats Working vation Society). Distribution y estado environmental education programs Group. The IUCNICat Specialist actual del jaguar (Panthera onca) en el that promote carnivore conserva- Group, in particular Peter Jackson Noroeste de Mexico. In Medellin, R.A., tion at the community level. BWG and Kristin Nowell, is thanked for its C. Chetkiewica, A. Rabinowitz, K.H. is initiating environmental educa- guidance and leadership to BWG. Reford, J.G. Robinson, E.W. Sanderson, y A. Taber (eds). El jaguar en el nuevo tion projects with both ranchers and Logistical support for field surveys milenio: Una evaluacion de su condicion school children in the border region was provided by Raymond Skiles and actual, historia natural y prioridades para and developing partnerships with Betty Alex at Big Bend National su conservacion. Fondo de Cultura, governmental agencies in the U.S. Park. Alison Gabel, Gerald Scoville, Economica, Wildlife Conservation So- ciety y UNAM, Mexico, D.F. and Mexico to raise awareness Dennis Sizemore, Kevin Crooks, and Tewes, M.E., S.E. Hayslette, and L.L. Laack. about these important species. The volunteers from Round River Con- 1997. Survivorship and mortality factors results of our future research find- servation Studies are thanked for con- of ocelots in South Texas. Caesar Kleberg ings will only enhance these long- ducting field surveys in Arizona and Wildlife Research Institute: Current Re- term partnerships, as additional in- New Mexico. The field participation search. formation becomes available about of Marcela Aranda, Dave and Nancy

132 Endangered Species UPDATE Vol. 18 No. 4 2001 Molecular Scatology as a Conservation tool Laura E. Farrell Museum of Comparative Zoology, Hatvard University. 26 Oxford Street, Cambridge, MA02138; IfarrellQoeb.hatvard.edu

Abstract The threat of predation on livestock by large carnivores represents a major impediment to the conservation of intact ecosystems throughout the world. Although it has taken nearly a century to learn the truth about predation on livestock in North America, advanced tech- nologies now exist to greatly expedite similar findings elsewhere. By examining the dietary ecology of puma and jaguar on a cattle ranch in the Venezuelan llanos, it is possible to help identify simple, effective methods to reduce livestock predation. In my study, dietary habits of the puma and jaguar were defined from feces (scats), kills, and ranch records. Scats were assigned to predator species through analysis of mitochondrial DNA from feces of wild carnivores. Based on dietary and ecological data of these two large carnivores, recommen- dations on livestock husbandry are made.

The puma (Puma concolor) and the than a decade it was the most accu- diet, and help lead to solutions for jaguar (Panthera onca) are large rate method for determining donor a range of conservation problems sympatric carnivores trying to sur- species. But bile acid assays can (Kohn and Wayne 1997). With the vive in increasingly fragmented give misleading results for closely use of scat-sniffing , non-bi- habitats of the Neotropics. Much related carnivores. The identity of ased recovery of samples from mul- of the range of these endangered puma and jaguar scats is based on tiple target species is now feasible animals has been converted to the presence of one bile acid that (Wasser et al. 1999), making popu- ranchlands, which now hold some appears in only 7 1% of jaguar scats, lation estimates for similar-sized of the last remaining natural habi- leaving a 29% possibility of mis- sympatric species possible. By ana- tats in Latin America. In much of taking jaguars for pumas (Taber et lyzing the prey content of scats the Neotropics, the major cause of al. 1997). Nevertheless, diameter from undisturbed habitats, we can mortality for large felids is perse- of puma and jaguar feces in Taber's determine the preferred prey of car- cution by cattle ranchers for alleged study overlapped in almost all size nivores. If we manage this prey ap- predation on livestock. To conserve ranges. The need for a more de- propriately, attacks on livestock by viable populations of large cats it finitive analysis is apparent. hungry carnivores can be reduced. is necessary to find some resolution The advent of polymerase chain The molecular method is com- to this problem. This study at- reaction (PCR) has enabled scien- pletely noninvasive, though remov- tempts to accurately identify the tists to acquire information through ing all scats may confuse animals predator species involved in live- molecular analysis. Hoss et al: by removing territorial markers, stock depredation, and to provide a (1992) used genetic techniques to which also signal reproductive sta- clearer understanding of carnivore analyze mitochondrial DNA tus (Brown et al. 1994). The field interactions with livestock. (mtDNA) from sloughed off colon collection methods for this study Dietary habits can be accurately cells found in wild scat in order to were simple and effective; samples determined noninvasively only by identify Italian bears. By isolating were collected, air-dried, and then a thorough analysis of scats-the DNA from scats and checking it stored at room temperature. Using end product of predation. Unfor- against reference samples, it is now these collection and storage meth- tunately, accurate analysis has possible to accurately determine ods, a recent lab run produced posi- eluded scientists until recently. which species, and sometimes even tive identifications from scats that Scats have traditionally been sorted which individual left the sample were seven to ten years old (in to donor species by diameter. In (Kohn et al. 1995; Kohn et al. 1999; preparation). Samples were trans- 1980 thin layer chromatography Ernst et al. 2000). ported without freezing or excess analysis of bile acids was intro- Molecular scatology can pro- bulk, hazardous chemicals, or cum- duced (Major et al.), and for more vide new insights into the carnivore bersome permit restrictions. Scats

Vol. 18 No. 4 2001 Endangered Species UPDATE 133 are the only item from Appendix I Jaguar species exempt from CITES con- > trols (Gerloff et al. 1995), although Puma other restrictions, such as those imposed by the USDA, may apply. Fox - Carnivore scats were collected Ocelot - opportunistically from roads and trails on Hato Piiiero, a cattle ranch in the llanos, the seasonally flood- 0 10 20 30 40 50 60 70 80 90 100 110 120 ing savanna of western Venezuela. Body weight (kg) Samples were dried and a small portion of each scat was stored for genetic analysis (Farrell et al. 2000). Samples were then analyzed Linares 1998 for prey contents; remains were vi- Figure 1. Overlap in body size of the four main carnivores on Hato Piiiero, sually identified to the lowest taxon Venezuela (Linares 1998). possible by examining teeth, claws, bones, fur, feathers, and scales. For this study, all less than large predator categories-first to find a cytochrome b sequence one kilogram were considered based on size of scat (traditional with the closest match. When that small-mostly rodents, but includ- method) and then based on DNA came back as a fruitfly (Drosophila ing small marsupial opos- identification. melanogaster; likely apprehended sums (Marmosa sp.). Medium For the preliminary analysis by while the scat was collected), prim- mammal prey ranged from 1 to 15 scat size, scats were split into large, ers that targeted carnivores were kilograms, including armadillo assumed to be puma and jaguar, and designed. The sample was (Dasypus novemcinctus), , sloth small, assumed to be ocelot and fox. reamplified with these new prim- (Bradypus variegatus), opossum (Di- Based on previous studies (L. ers and found to be a perfect match delphis marsupialis), juvenile pec- Emmons, personal communication; with our ocelot reference. Target- cary, and newborn calves (Bus bus). Fernandez et al. 1997), 25 mm dry ing a shortened fragment of 147 Large mammals were anything over diameter was chosen as an arbitrary base pairs also increased the per- 15 kilograms, such as adult capybara breaking point large enough so that centage of successful samples, be- (Hydrochaeris hydrochaeris) and it was beyond the range of most fox cause fecal DNA is often degraded. peccary, deer, giant anteater and ocelots. Twenty of 34 samples (59%) were (Myrmecophaga tridactyla), and ju- For the molecular analysis, a successfully sequenced. venile and adult cows. portion of the mitochondria1 cyto- Three jaguar and two fox In the llanos, the ocelot chrome b gene was chosen as the scats were identical to control (Leopardus pardalis) and crab-eat- DNA marker to assign unknown samples, as were five puma scats ing fox (Cerdocyon thous) each 'scats to predator species. Samples (one puma scat differed by one average about six to 10 kg in body were washed, and DNA extracted base pair). The sequences from weight, the puma about 40 to 50 kg, from the isolate and amplified us- 10 ocelots showed more varia- and the jaguar up to 100 kg (Figure ing PCR (Farrell et al. 2000). The tion, which makes sense in light of I). But, body size is not necessar- resulting strands of DNA were se- a study by Eizerick et al. (1998) ily indicative of scat size. Puma quenced and compared to samples showing a relatively high degree of scats have been identified down to extracted from blood of the carni- intraspecific variation among the 19mm in diameter, and ocelot scats vore species at the study site. Even smaller Neotropical felids. up to 27 mm. Because of this over- this test wasn't perfect at first-one Puma, ocelot, and fox scat sizes lap, it was necessary to define prey sample that could not be matched overlap in a narrow range around contents of almost all size predator to a known carnivore was compared 25mm diameter, and puma and jag- scats to accurately define the di- to sequences posted in GenBank us- uar overlap at larger diameters be- etary habits of puma and jaguar. ing the BLAST program (National tween 32 and 37 mm (Figure 2). Frequency of occurrence of prey Center for Biotechnology Informa- One individual can leave scats of types were separated into small and tion (NCBI), www.ncbi.nlm.nih.gov) various sizes, and the range of body

134 Endangered Species UPDATE Vol. 18 No. 4 2001 4 Scats to species using mtDNA be highly significant (X- = 7.36, p<0.0 1, ldf). With differentiation by size, 83% of ocelot and fox scats were misclassified as large preda- n 3 Jaguar tor scats, and 12% of puma and jag- uar scats were misclassified as n = 5 Puma small predator scats. These find- ings are preliminary and the sample Fox n = 2 - size is small, dietary results may n = 10 Ocelot - change when scats from the last two years of the three year project are analyzed. 19 20 25 30 35 40 41 With DNA analysis, predators Scat diameter (mm) are discernable to the species level Figure 2. Overlap in scat sizes of the four main carnivores when the scat (Table 2)- Thebbmass of each Prey identity is confirmed by molecular analysis. type is then calculated to learn its relative importance in each carnivore's diet (Farrell et al. 2000). Dietary resources are divided more between large and small carnivores sizes can vary greatly within spe- cies. For example, male jaguars in the llanos average about 100 kilo- grams, whereas males in the Cen- Table 1. A comparison of prey frequencies with scats separated by size and molecular analysis. One jaguar scat was disqualified from tral American rain forest average 55 the dietary analysis for containing bait (from Farrell et al. 2000). Small kg. predators = Fox and ocelot. Large predators = Puma and jaguar. Using size to define dietary analyses, smaller carnivores appear Frequency (%) of Prey Types to ingest solely mammalian prey Scat size mtDNA (Table 1, left side). They even ap- Small Large pear to take large prey, though the Small Large predators Predators thought of an ocelot or fox subdu- Predators Predators ing and killing an 18 kg collared 525mm 125mm peccary (Tayassu tajacu) may seem Small unusual. Large predators appear to 75 46 64 take prey from all categories, pre- mammals dominantly small mammals. Medium DNA analysis of samples 12.5 12 3 50 mammals shows that almost the inverse is true. Large carnivores prey exclu- Large 12.5 9 50 sively on medium and large mam- mammals mals, and small carnivores take a wide range of prey, including ev- Reptiles 9 9 erything except large mammals (Table 1, right side). Analysis of one Birds 12 12 scat with a size indicative of a small ocelot (19.25mm), clarified that it Fish 3 3 was a puma who ate the peccary. McNemar's test (Harrison Crabs 9 9 1996) showed the difference be- tween identification of scat donor n=3 n=16 n=12 n=7 by size and by genetic analysis to

Vol. 18 No. 4 2001 Endangered Species UPDATE 135 Table 2. Frequency of prey biomass with predator scats identified ducive to predation. Seven verifi- using molecular analysis (from Farrell 1999). able incidences of livestock preda- Jaguar Puma Ocelot Fox tion were examined, and six of n=2 n=5 n=10 n = 2 these confirmed as pumas (Farrell 1999). Frequency and location of predation by cats on cattle was de- termined from 10 years of ranch records that reported causes of live- stock mortality (Table 3: Hato Large 72 54 mammals Pifiero, unpublished data). When Reptiles 43 observations of livestock killed during my study are combined with Birds evidence from ranch records and Fish molecular data, the preference for different sizes of prey between the Crabs 12 two cats becomes clearer. Jaguars Iselect for very large adult prey. They took only a very small per- Table 3. Predation on cattle (calves and adults) by puma and jaguar on Hato Piiiero from January 1987 through Oc- centage of the total cattle lost (el % tober 1996, as reported by ranch workers (Hato PiAero, un- according to ranch records), yet published data; from Farrell1999). *There was found to be these cows were typically full a discrepancy of 27% between causes of livestock mortal- grown and a greater financial loss ity reported by ranch workers and evidence at the site when per head. As the more endangered reported incidences of predation were investigated in 1999. of the two felids, perhaps jaguars should be allowed to keep practic- Total Puma Jaguar ing business as usual, and a com- pensation program for ranchers be Number of mortalities attributed 310 275 35 initiated. Pumas are pinpointed as to predation* a greater problem; they are more general in prey selection and con- % Predation 89 11 centrate on medium and large size prey-often in the form of newborn

O/O total cattle mortalities from and juvenile calves. Their prefer- of 8,4 7.5 <, all causes (n=3672) ence for calves and juveniles when preying on livestock corroborates data from genetic identification of puma scats that revealed no prey than between same size sympatric missing tag-along dog had been item larger than a juvenile calf. carnivores; puma and jaguar versus consumed by a puma. The samples While jaguars prefer moister fox and ocelot. Jaguar scats con- containing dog and pig bait illus- habitats of lower elevations; the pu- tained a peccary and a cottontail trate that this method can be used mas preference for dry habitat types rabbit (Sylvilagus floridanus). A to pinpoint problem predators. is significant. Pumas move to third was disqualified from the di- Dietary data derived from the dryer, nonflooding ranges for the etary analysis because it contained genetic analysis, along with infor- wet season. Unfortunately cattle a wild pig set as bait while trapping mation from kills recovered during are also moved to these areas at the cats for radio-telemetry. Five puma this study and ranch records, can of- same time. The birthing of insemi- scats contained a couple of juvenile fer suggestions to help decrease nated cows (each representing an peccaries, an armadillo, one deer livestock predation. Livestock and investment of over $100 US) is (Odocoileus virginianus), and a do- wild kills were examined when lo- timed as well for the beginning of mestic guard dog (Canis cated (n=15) and habitat conditions the wet season in pastures along familiarus), confirming the suspi- were recorded to discern where cats the edges of the ranch. In these cion of ranch workers that their kill, and what conditions are con- areas humans harvest much of the

136 Endangered Species UPDATE Vol. 18 No. 4 2001 natural prey, so it is no surprise that puma in the Neotropics. and jaguar feces through bile acid chro- these pastures exhibit the greatest matography. Journal of Wildlife Manage- ment 61506-5 10. incidences of livestock 'predation. Acknowledgements Gerloff U, C. Schlotterer, K. Rassmann, I. Pumas arriving thin and hungry from I would like to thank Sam Wasser Rambold, G. Hohmann, B. Fruth and D. desolate ranges at the end of the dry and Chris Clarke, who generously Tautz. 1995. AmpWcation of hypervariable season are greeted by a landscape allowed me to learn in the Center simple sequence repeats (rnicrosatellites) flush with naive, newborn prey that for Wildlife Conservation's molecu- from excremental DNA of wild living bonobos (Panpaniscus). Molecular Ecol- are just learning to walk. lar lab, Joe Roman who helped with ogy 4515-5 18. It is important to incorporate many facets of this study, Brian Harrison, J.M. 1996. Categorical Data Analy- different types of data and the views Bowen who encouraged this along, sis. Jay M. Harrison, IFAS Dept. of Statis- of different parties to create solu- and Me1 Sunquist, without whom tics. University of Florida, Gainesville, FL. Hoss M., M. Kohn, S. PiCibo, F. Knauer and tions that will be effective in help- this would not have been possible. W. Schriider. 1992. Excrement analysis by ing both predators and local ranch- Martin Smith reviewed earlier ver- PCR. Nature 359: 199. ers. This study verified that pumas sions of the manuscript and his sug- Kohn M., F. Knauer, A. Stoffella, W. are a greater problem than jaguars gestions improved the paper. Many Schroder and S. Paabo. 1995. Conserva- on Hato Piiiero. By determining others helped as well, special tion genetics of the European brown bear-a study using excremental PCR of which prey carnivores favor thanks to the llaneros on Hato nuclear and mitochondrial sequences. through scat analysis, management Pinero. This study was part of work Molecular Ecology 4:95-103. of the preferred prey can help re- done for a Masters thesis at the Uni- K0hnM.H. andR.K. Wayne. 1997. Facts from duce predation problems. To test versity of Florida, Gainesville, with feces revisited. Trends in Ecology and Evo- lution 12:223-227. deterrence, electric fencing was Dr. Me1 Sunquist, who put together Kohn M.H., E.C. York, D.A. Kamradt, G. used in one of the hardest hit ma- the field project. Haught, R.M. Sauvajot and R.K. Wayne. ternity pastures during a later phase 1999. Estimating population size by of this study, and found to curtail Literature cited genotyping faeces. Proceedings of the Royal predation by 100% (Scognamillo et Brown, J.L., S.K Wasser, D.E. Wddt and L.H Society of London B 266:657-663. Linares, O.J. 1998. Mamfferos de Venezuela. al. 1999). One noteworthy finding Graham. 1994. Comparative aspects of ste- roid hormone metabolism and ovarian ac- Sociedad Conservationista Audubon de Ven- of my study is that when water buf- tivity in felids, measured non-invasively in ezuela, Caracas. falo (Bubalis bubalis) were placed feces. Biology of Reproduction 51:776-786. Major M., M.K. Johnson, W.S. Davis and T.F. with cattle, predation mortalities in Eizirick E., S.L. Bonato, W.E. Johnson, P.G. Kellog. 1980. Identifying scats by recov- Crawshaw Jr., J.C. Vi6, D.M. Brousset, S.J. ery of bile acids. Journal of Wildlife Man- one long-suffering maternity pas- agement 44290-293, ture decreased close to zero O'Brien, F.M Salzano. 1998. Phylogeographic patterns and evolution of Scognamillo, D., I.E. Maxit, M. Sunquist, (Andres Rodriguez, personal com- the mitochondrial DNA control region in two and L. Farrell. In press. Ecologia del jag- munication). Water buffalo are al- Neotropical cats (Mammalia, Felidae). Jow- uar y el problema de la depredacion sobre ready being raised on this and other nal of Molecular Evolution 47613-624. ganado en Hato Piiiero, Venezuela. Manu- script submitted for publication in El Jag- ranches throughout the llanos for Ernest HB, M.C.T. Penedo, B.P. May, M. Syvanen and W.M. Boyce. 2000. Molecu- uar en el nuevo milenio. Una evaluacion meat and cheese and mix well with lar tracking of mountain lions in theyosemite de su estado, deteccion de prioridades y cattle, while apparently affording Valley region in California: genetic analysis recomendaciones para la conservacion de them some protection from preda- using microsatellites and faecal DNA. Mo- 10s jaguares en America. Medellin, R. A., lecular Ecology 9:433-441. C. Chetkiewicz, A. Rabinowitz, K. H. tion (Farrell 1998). Redford, J. G. Robinson, E.W. Sanderson, Vast portions of puma and jag- Farrell L.E. 1998. Water buffalo as a conser- vation tool. Water Buffalo Newsletter y A. Taber, eds. Universidad Nacional uar ranges are dedicated to livestock 10(4):2 The American Water BuffaloAsso- Autonoma de Mexico~WildlifeConsema- production-often on ranches of ciation, Gainesville, FL. tion Society. Mexico D. F. 80,000 to 120,000 hectares that con- Farrell L.E. 1999. The ecology of the puma Taber A.B., A.J. Novaro, N. Neris and F.H. Colman. 1997. The food habits of sympat- tain large areas of natural habitat. and the jaguar in the Venezuelan llanos. MS thesis. University of Florida, Gainesville. ric jaguar and puma in the Paraguayan Owners who are confident that their Farrell, L.E., J. Roman and M.E. Sunquist. Chaco. Biotropica 29:204-213. stock is safe are less likely to go hunt- 2000. Dietary separation of sympatric car- Wasser S., M. Parker and B. Davenport. 1999. ing for these elusive cats. Reducing nivores identified by molecular analysis of Noninvasive DNA sampling using scat sniff- the threat of felid predation on live- scats. Molecular Ecology 9: 1583-1590. ing dogs. Society for Conservation Biology Abstracts. Thirteenth Annual Meeting, Col- stock will have broad implications for Fernandez G.J., J.C. Corley and A.F. Capurro. 1997. Identification of cougar lege Park, Maryland. future conservation of jaguar and

Vol. 18 No. 4 2001 Endangered Species UPDATE 137 Linking Snow Leopard Conservation and People-Wildlife Conflict Resolution: Grassroots Measures to Protect the Endangered Snow Leopard from Herder Retribution

Rodney Jackson Snow Leopard Conservancy, 236 North Santa Cruz Ave., Suite 201, Lost Gatos, CA95030, USA; (408) 354-6459; (fax) (408) 654-5869; [email protected] Rinchen Wangchuk Field Program Director, Snow Leopard Conservancy, Leh, Ladakh, India

Abstract Livestock depredation has become a signijicant problem across the snow leopard's (Panthera uncia) range in Central Asia, being most severe in and near protected areas. Such predation, especially incidents of "surplus killing," in whichfive to 100 or more sheep and goats are lost in a single night, almost inevitably leads herders to retaliate by killing rare or endangered carnivores like snow leopard, WOK and lynx. Ironically, such loss can be avoided by making the night-time enclosures predator-prooj improving animal husbandry techniques, educating herders on wildlife consewmion and the importance of protecting the natural prey base, and by providing economic incentives like handicrafts skills training and marketing, along with carefully planned ecotourism trekking and guiding. The author explores innovative conservation initiatives in the Himalaya (Ladakh and Tibet) and Mongolia, which also build local capacity, self-reliance, and stewardship for nature using Appreciative Participatory Planning and Action, or APPA, techniques. The most sound conservation investments are those contingent upon establishing direct linkages with biodiversity protection, ensuring co-financing and reciprocal responsibility for project activities, encouraging the full participation of all stakeholders, and assuring regular monitoring and evaluation of the village-based agreements (embodied in Action Plans).

Introduction A similar situation in Hemis ness to co-exist with predators Wildlife damage is a major source National Park, Ladakh, Jammu and hinges upon reducing depredation for conflict between local com- Kashmir State, led to the establish- to an acceptable level while im- munities and protected areas ment of a compensation scheme, proving incomes to help offset un- managers in the Himalaya but within two years the sponsor- avoidable losses of livestock. (Kharel 1997; Jackson et al. ing Ladakh Wildlife Department This field note describes grass- 1996; Oli et al. 1994). In India's found itself committing 60% of its roots initiatives being undertaken in Kibber Wildlife Sanctuary, Mishra annual $26,000 budget to the pro- Hemis National Park to alleviate (1997) noted that 18% of livestock gram. Payment takes up to two livestock loss to predation and to holdings were killed by snow leop- years, with claimants being paid encourage herders to become effec- ard (Uncia uncia) and wolf (Canis only 10 to 30% of their animal's tive stewards of the snow leopard, lupus) for an estimated total value market value. Understandably re- its prey and its habitat. of US $128 per household per an- lations between local people and num-a very significant economic the park have deteriorated, with re- Community-based conflict impact given per annual cash in- taliatory killing constituting a ma- alleviation initiatives in comes of $200 to $400. Villagers jor threat to both snow leopard and Hemis National Park claimed predation rates increased wolf. Because local livelihoods are Established in 1981, this park cov- after sanctuary establishment, intimately bound with long-stand- ers 3,350 square kilometers in the while surveys indicated dramatic ing patterns of agro-pastoralism, re- TransHimalayan Range (Fox and increases in livestock numbers ac- location of people or the exclusion Nurbu 1990). Besides offering ex- companying changes in animal hus- of livestock is not a feasible solu- cellent snow leopard habitat, the bandry systems (Mishra 2000). tion. Rather, local people's willing- park's four species of wild sheep

138 Endangered Species UPDATE Vol. 18 No. 4 2001 and goats give it international numbers natural prey and biomass in predator densities, habitat suit- biodiversity importance. About only invites loss to wild predators. ability and herding patterns. 1,600 people live in 16 small settle- Historically there has been better The household survey was fol- ments scattered in three valleys. emphasis on daytime guarding, and lowed by a workshop held in They grow barley and a few veg- problem predators were controlled Markha village in association with etables, and own more than 4,000 through trapping and other tradi- the Ladakh Wildlife Department, head of livestock, of which 8 1% are tional control methods. With more national and international non-gov- sheep and goats, and 1 1% are yaks, children going to school and youths ernmental agencies (NGOs). The cattle and crossbreeds. Tourism reticent to assume this hard liveli- primary objectives were to (1) iden- provides an important source of hood, even highly vulnerable small- tify cost-effective and ecologically supplementary income. Ladakh bodied livestock are left to graze compatible measures for reducing was opened to tourism in 1974, and unattended. While baseline docu- livestock losses; (2) Train park the Markha Valley circuit through mentation is lacking, predator num- staff, NGOs and villagers in wild- Hemis National Park remains the bers appear to have increased due life damage alleviation techniques; most popular trekking route, with to park regulations prohibiting and (3) Promote community-based about 5,000 visitors per year. hunting and patrolling by park wildlife stewardship and enhance We surveyed 79 households liv- guards. As Figure 1 suggests, dep- awareness of the opportunities to ing within or immediately adjacent redation rates vary with locality, conserve even "problem" species. to the park to determine livestock presumably reflecting differences Using a highly participatory ownership patterns, document dep- redation losses and map the depre- dation-prone areas or "hotspots" (Bhatnagar et al. 1999). Over half the households interviewed lost one to 15% or more of their domestic stock to predators, or 492 animals valued at USD $23,500 over a 14- month period. Snow leopard and wolf were associated with 55% and 3 1% of the presumed depredation incidents respectively, with sheep and goats constituting 75% of the stock lost, followed by yak-cattle (13%) and horses (8%). Three settlements incurred 54% of the depredation. Losses incurred from snow leopards entering poorly con- structed corrals accounted for 14% of all incidents (N=210), but nearly 50% of all livestock lost-under- standably arousing considerable anger among the livestock owners. Along with poorly constructed livestock pens, investigations into the root causes of depredation im- plicated lax daytime livestock guarding practices. Stock was al- lowed to forage in areas with well- broken terrain and cliffs, prime habitat for snow leopard (Jackson et al. 1996). The fact that domes- Snow leopard (Uncia uncia). Photograph by M. Elsbeth tic livestock now substantially out- McP hee

Vol. 18 No. 4 2001 Endangered Species UPDATE 139 process known as Appreciative Par- ticipatory Planning and Action (APPA), workshop participants and villagers examined root causes of depredation and identified a series of measures aimed at reducing dep- redation loss, improving household incomes and promoting wildlife conservation. APPA combines con- cepts from Appreciative Inquiry (used in business leadership train- ing) and Participatory Learning and Action (PLA, Pretty et al. 1995), in a collective inquiry and planning process aimed at fostering effective group action. It operates under two complimentary premises: (1) What Figure 1. Depredation hotspots you seek is what you find-"if you look for problems, then you will find more problems" or conversely, "if you look for successes, you will fined conservation objectives. remedial actions that were environ- find more successes;" and (2) What 2) Participation-the active mentally responsible (i.e., compli- you believe is what matters most- and equitable involvement of each ant with park regulations and spe- "if you have faith in your vision or stakeholder group is promoted cieslhabitat management require- ideas for the future, and if these are throughout the project to ensure all ments); economically sustainable believable, you can achieve success affected households will be benefit- within the local context; socially without waiting for government or ted and to encourage participation responsible (e.g. building upon outside agents to take you there." irrespective of gender, age or eco- proven traditions and cultural val- APPA is practiced through an nomic status. ues which protect nature); and iterative process that seeks to (1) 3) Reciprocity-All stake- which are implemented under a discover the community's strengths holders, whether outside donor, mutually agreed-to (and signed) and its valued resources; (2) envi- local NGO, government, or vil- work-plan that sets forth the re- sion short-term and long-term fu- lagers are expected to make a re- sponsibilities, contributions and tures if resources were mobilized ciprocal contribution within their obligations of each partner. and the community acted in con- means (e.g., cash, materials, la- Markha villagers concluded cert; (3) design a basic action plan bor, or in-kind service). that their best option lay in replac- for guiding development and nature 4) Responsibility-The ben- ing the existing four winter corrals protection in ways that substan- eficiary community must be will- with three larger predator-proof tially limit long-term dependency ing to assume responsibility for structures placed side-by-side and upon outside financial sources or meeting the conservation objectives sharing inner walls. They donated technical "know-how;" and (4) mo- and for maintaining any their labor and provided on-site ma- tivate participants to initiate com- infrastructural development. There terials (stones and mud), while ex- munity-improvement actions imme- should be clear penalties for infringe- ternal donors provided off-site ma- diately, and largely on their own. ment by any of the participants. terials (wire mesh, roofing poles, Outside donor support was 5) Monitoring-Stakeholders and secure doors). Construction only offered if the following pro- should employ simple but realistic was scheduled for spring, but was visions were met: indicators for monitoring project delayed due to frozen ground. Also, 1) Conservation-Biodiversity impact and performance, described corrals had to be 15 feet longer than conservation is the primary motiva- in the Action Plan prepared jointly the plans indicated because the vil- tion behind external investment, and and signed by the key parties. lagers deliberately underestimated therefore all project activities must External expertise is blended their livestock holdings, fearing be implicitly linked with clearly de- with local knowledge in designing they would be taxed more by the

140 Endangered Species UPDATE Vol. 18 No. 4 2001 government for reporting actual being offered skills training to en- their commitment to finding long- herd sizes. Unfortunately, they used hance their summer tea-house op- term solutions to this vexing problem. the corral before it was fully preda- erations by improving menus, en- Literature cited tor-proofed, and lost 29 animals to suring hygienic conditions, and Bhatnagar, Y.V., R. Wangchuk, and R. a snow leopard. The outside donors building campground facilities. A Jackson. 1999. A survey of depredation and related wildlife-human conflicts in felt some responsibility for the loss key next-step will be to use the Hemis National Park, Ladakh, Jammu and called a community meeting. "parachute cafes" or tea-houses as and Kashmir, India. Unpublished Re- The villagers, however, assumed focal points for providing tourists port, International Snow Leopard Trust. full responsibility for what had hap- and local communities with wild- 20 pages. Fox, J.L. and C. Nurbu. 1990. Hemis, a Na- pened. Their reasoning was as fol- life conservation education. tional park for Snow Leopard in India's lows: there had been a death in one We believe this approach is Trans-Himalaya. International Pedigree of the families, just before the depre- highly effective in mobilizing ru- Book of Snow Leopards 6:71-84. dation. As the other six affected ral communities toward greater Jackson, R.M., G. Ahlborn, M. Gurung, households had only lost one or two self-reliance and thus a more har- and S. Ale. 1996. Reducing livestock depredation losses in the Nepalese animals, they all agreed that a tradi- monious long-term relationship Himalaya. Proceedings Vertebrate Pest tional Mountain Spirit had been re- with the National Park in which Conference 17:241-247. sponsible for the snow leopard's visit. they live, and on whose resources Kharel, F.R. 1997. Agricultural crop and We believe that by predator-proofing they depend so heavily. APPA livestock depredation by wildlife in a village's corrals we are removing builds pride by highlighting posi- Langtang National Park, Nepal. Moun- tain Research and Development as many as five to 10 snow leopards tive community attributes and 17(2):127-134. from risk of retaliatory killing. building upon traditional values and Mishra, C. 1997. Livestock depredation by successes. NGOs are the most ob- large carnivores in the Indian trans- Conclusions vious vehicle for facilitating com- Himalaya: conflict perceptions and con- servation prospects. Environmental Con- At the broader level, the future of munity-based integration of conser- servation 24(4):338-343. these protected areas hinges on the vation and development; however, Mishra, C. 2000. Socioeconomic transition degree to which the basic concerns, the sponsoring agency must be will- and wildlife conservation in the Indian needs and aspirations of the local ing to make a long-term commit- Trans-Himalaya. Journal Bombay Natu- people are addressed. Over the ment to their rural stakeholders ral History Society 97(1):25-32. Oli, M.K., I.R. Taylor, and M.E. Rogers. long-term, the most cost-effective (Sanjayan et al. 1997). 1994. Snow leopard (Panthera uncia) approach for cash-strapped devel- predation of livestock - an assessment oping countries may lie in promot- Acknowledgments of local perceptions in the Annapurna ing a set of carefully designed and This work was initiated in the senior Conservation Area, Nepal. Biological Conservation 68(1):63-68. monitored community-based stew- author's capacity as Conservation Pretty, J.N., I. Guijt, I. Scoones, and J. Th- ardship initiatives in which local Director of the International Snow ompson. 1995. A Trainer's Guide for Par- people benefit from the presence of Leopard Trust, now being continued ticipatory Learning and Action. IIED wildlife, including predators. under the Snow Leopard Conser- Participatory Methodology Series, Inter- While our initial effort focused on vancy. Special thanks to the J&K national Institute for Environment and Development, London. 267 pages. reducing loss of livestock to preda- Wildlife Department and LEDeG for Sanjayan, M.A., S. Shen, and M. Jansen. tors, we are now concentrating on co-sponsoring the workshop in 1997. Experiences with integrated-con- measures aimed at helping local Markha. The senior author is in- servation development projects in Asia. people capture more benefits from debted to Rinchen Wangchuk and the World Bank Technical Paper 388:l-41. tourism. For example, women are residents of Hernis National Park for

Vol. 18 No, 4 2001 Endangered Specles UPDATE 14 1 Canid Conservation The Status of the Wolf Population in Post-Soviet Kyrgyzstan CJ Hazel1 State University of New York, Environment Science and Forestry, 133 lllick Hall, 1 Forestry Drive, Syracuse, NY 13210; [email protected]

Abstract The countries that made up the former Soviet Union are home to the largest existing wild gray wolf(Canis lupus) population. This makes the status of wolves in these areas of particular importance. Kyrgyzstan is a former Soviet republic located in Central Asia that became indepen- dent in 1991. Russia and other former Soviet republics with wolves reported in 1999 that their wolfpopulations were either stable or increasing. Kyrgyz oficials also consider their wolf populations to be increasing since independence. At the same time, oficials report that popula- tions of deer, wild boar, and other wildlife are decreasing drastically. Between January and May 1999, I made sevenfield trips in southern Kyrgyzstan to look for evidence to support oficial reports. I visited two zapovedniks (strictly protected areas), one national park and two national forests (less protected), and two rural areas with no wildlife protection to look forfield signs and to collect anecdotal informationfvom local villagers. I found abundant wolf sign at only two sites and abundant deer sign at only one site. Villagers tended to say that there were many wolves, but they were 20 km away. They also said that deel; boar, and other wildlife were dificult tojnd now. This evidence supported oflcial reports that wildlife populations were decreasing but contradicted reports that wolfpopulations were increasing. Government records show that most species of game animals and sheep, the primary livestock animal, have decreased by close to 50% in the south since independence. Data also showed that wolfpopulations in the south have dropped by 43%. This evidence suggests that oficial reports that wolfpopulations in Kyrgyzstan are stable or increasing are inaccurate and wolfpopulations may actually be declining.

Introduction In 1999, Wolf International, the sessment of the impact these Gray wolves (Canis lupus) used to magazine of the International Wolf changes are having on wolf popu- roam over most of the Northern Herni- Center, reported on the status of lations is important. sphere. Human persecution over the wolf populations worldwide, in- The Kyrgyz Republic is a former last few centuries has resulted in very cluding some areas of the former Soviet republic in Central Asia that small, isolated populations or extirpa- Soviet Union. Of the Common- became independent in 199 1. It is a tion in much of North America and wealth of Independent States (CIS) small country of just under 200,000 Western Europe. The area encom- countries reporting, most list their krn2 with a population of about 4.6 passed by the former Soviet Union is populations as increasing or stable million people. Over 90% of the the home of the largest remaining popu- with wolves being unprotected or terrain is mountainous with eleva- lations of gray wolves (Theberge considered game species (Route tions ranging up to 7,439 meters. 1991). The Soviets have always con- and Aylsworth 1999). Since the Almost 40% of the land area is con- sidered wolves to be a pest species and break up of the Soviet Union from sidered to be uninhabitable due to have traditionally supported strong, 1989 to 1991, most of that region elevation, glaciers, or the presence government-sponsored wolf control has been experiencing severe eco- of bare rock. Less than 4% of the programs. Historically, during eco- nomic decline (Anderson and land area is currently forested. It has nomic downturns, wolf populations Pomfret 2000). Given the changes an arid, alpine climate and much of have increased due to less money being currently taking place, both po- its economy is agriculturally based available for wolf control (Bibikov 1993). litical and economic, an as- with livestock grazing, particularly

142 Endangered Species UPDATE Vol. 18 No, 4 2001 ests, and alpine meadows at eleva- tions ranging from 2,000 to 3,000 meters. The major commonality for these areas was that all of them were historically rich in wildlife. I also collected anecdotal information from local people on wolves and the abun- dance of deer and other wildlife. In particular, I asked about changes they had noticed since Kyrgyzstan became independent. In addition, I looked at government records from the Hunt- ing Institution and the Agricultural Ministry of Osh Oblast (province) in southern Kyrgyzstan to determine the official estimates of both wildlife and livestock populations and the changes from pre- to post-independence. Commonwealth of Independent States (CIS): Central Asian States Results Out of the seven locations surveyed, there was only one that had abundant sheep, a primary activity (Ministry of From January to May 1999, I signs of both wolves and potential Environmental Protection 1998). conducted a study to determine if the wild prey species. This was in one perception that wolves were increas- of the rural, unprotected areas. Wolf Evaluating the current status of ing in abundance in spite of decreas- tracks were abundant both on the wolf populations ing wildlife populations could be sub- mountain ridges and down into the Information on the condition of wild- stantiated. I conducted seven field river valley near the villages. Roe life in Kyrgyzstan that was collected surveys in the southern part of the deer (Capreolus capreolus) were also as part of a biodiversity assessment country by going out with local abundant in the area, though there completed in 1999 indicated that, in guides to look for wolf signs as well were no signs of wild boar (Sus general, wildlife populations were as signs of potential wild prey spe- scrofa), which used to be common in declining (Ministry of Environmen- cies. These surveys were not meant the area. We did locate one deer kill tal Protection 1998). In contrast, the to be exhaustive or systematic. I site about one km from a village and general perception on the part of the asked the guides to show me the most also a marking post with scat on a public and government officials was likely places that wolf sign would be mountain ridge 12 km from the vil- that wolf populations were actually found. I hoped to survey the areas lage. The scat consisted entirely of increasing. People in villages were that were more likely to have higher wool, indicating that these wolves had concerned about livestock losses to wolf populations than elsewhere near been eating sheep. wolf depredation. The perception the village. This way, the results were The other unprotected rural lo- that wolves were an increasing threat more likely to indicate an abundance cation also had abundant signs of to the economic well being of villag- of wolves and result in a conserva- wolves coming down from the moun- ers contributed to the government tive measure of possible declines. tain ridges into the river valley near policy of paying a substantial bounty The surveys included two the villages. The only wildlife spe- for killing wolves. The bounty in zapovedniks (protected areas with no cies that appeared abundant in the 1998 was 1000 som ($20) for an adult recreational use or hunting allowed), river valley, however, was red mar- male and 1500 som ($30) for an adult two national forests and one national mot (Mamota caudata). There were female (Kurmanaliev, personal com- park (protected, but some uses al- no signs of deer or other larger mam- munication). This is a substantial lowed), and two rural areas with no mal species. The wildlife signs in the amount of money in a country where protected status. These areas also rep- other sites surveyed ranged from a over half the population makes less than resented a variety of ecosystem types few to no wolf signs with few signs 4,500 som a year (World Bank 1999). including juniper forests, walnut for- of other large mammal species.

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Vol. 18 No. 4 2001 Endangered Specles UPDATE 143 One of the zapovedniks, Sara Chilik, had some signs of wolves, deer, and wild boar, but the official wildlife counts indicated that the numbers of all were lower than they had been a decade earlier, prior to in- dependence. In Besh Aral Zapovednik, there were few signs of wolves in spite of high official esti- mates. There were no deer or boar signs and marmots were the only spe- cies that appeared abundant. There were no long-term population sur- veys available for Besh Aral so a Figure 1. Survey of roe deer populations in Osh Oblast (data from comparison of abundance from be- Osh Oblast Hunting Institution 1988-1999). fore to after independence was not possible. Anecdotal evidence indi- cated that in most areas, most wild- life species were declining in num- bers. Villagers were having to range further away when hunting to find game. Even wild boar, which had not been hunted much in the past due to the Muslim culture of the Kyrgyz, seemed to be in decline. In most of the villages near the field survey sites, the locals felt that wolves were still abundant, but that most of the wolves Figure 2. Survey of wild boar populations in Osh Oblast (data from were further away from the villages Osh Oblast Hunting Institution 1988-1999). than they had been before. Most cases of wolf depredation on live- stock that were related to me were from "the next village" rather than from the village I was in. wild boar in Osh Oblast from 1988 for wolves, I also examined the agri- to 1999 (Osh Oblast Hunting Institu- cultural census data for Osh Oblast Official estimates tion 1988- 1999). These estimates from 1987 to 1997 (Osh Oblast Sub- The Government Hunting Institu- showed a substantial decrease in the sistence Farm Administration 1987- tions of Kyrgyzstan are responsible number of deer and boar over this 1997). This was to determine if there for keeping yearly estimates of all time period. Deer populations de- had been changes in the total number games species and other species of clined by 39.4% while boar popula- of livestock that might be available interest throughout the country. tions declined by 55.7% from a few to wolves. According to the official While the numbers from the official years before the breakup of the So- count, the numbers of horses and surveys may not be accurate as to the viet Union to the present with the de- cattle have experienced only small actual population size for the animals clines starting around 1992, just af- declines over the 10-year period. surveyed, they are collected using the ter independence (Figures 1 and 2). However, sheep, which is the domi- same technique from year to year. These numbers coincided with the nant domestic species in the country, Therefore, while not relying on the results of the field surveys and with have steadily declined since indepen- numbers to represent true population the anecdotal information from both dence. There were almost two mil- sizes, I felt that they were still useful official and local sources. lion sheep in Osh Oblast before the to indicate trends. I compared the re- Since livestock plays such an irn- breakup of the Soviet Union and by sults of government estimates of the portant role in the Kyrgyz economy and 1997 the number had declined to less populations of wolves, roe deer, and can serve as an alternative prey base than 900,000-a drop of 55.6%.

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144 Endangered Specles UPDATE Vol. 18 No. 4 2001 cultural sector of the economy. This is significant because the economy is largely agriculturally-based (World Bank, 1999). The large state-owned collective farms of the Soviet Union have almost all been broken up and the assets distributed as part of the privatization process (Karimbiekov, personal communication). As a re- sult, most livestock is now privately held in small family farms with pri- vate herds being relatively sniall. The increase in poverty has also played a role in the overall decrease Figure 3. Survey of gray wolf populations in Osh Oblast (data from Osh Oblast Hunting Institution 1988-1999). of wildlife populations. More people are exploiting natural resources for their daily subsistence. Poaching has In addition to showing declining with the breakup of the Soviet Union increased significantly. Habitat de- populations for wild prey species, the and the coming of independence. struction is also an increasing prob- Government Hunting Institution sur- Since the breakup of the Soviet lem with the need to use natural re- vey data also showed a marked de- Union, Kyrgyzstan, like most of the sources for both food and fuel due to cline in wolf abundance from ap- former republics, has been experienc- a lack of alternatives that were pre- proximately 1100 wolves in Osh ing a severe economic decline. Over- viously available. In addition, the Oblast to just over 600 -a decline all the country has experienced declines changes in the structure of livestock of 44.3% over the same time period in productivity and GDP, resulting in a herds has brought about a change in (Figure 3) (Osh Oblast Hunting In- sharp increase in the unemployment grazing patterns, resulting in in- stitution 1988-1999). This would rate (World Bank 1999). During the creased competition for resources be- seem to indicate that even the survey early years of independence, there was tween wild and domestic species information collected by the govern- a serious decline in income for most (Ministry of Environmental Protec- ment does not support the official people (Anderson and Pomfret 2000). tion 1998). The reason that this eco- stance that wolf populations have in- Pensions are no longer sufficient for the nomic decline hasn't resulted in an in- creased substantially since indepen- elderly to live on,and even people who crease in wolf populations is that the dence. The field survey results, as did have jobs frequently were not paid nature of the economy itself has well as much of the anecdotal infor- for months at a time (personal obser- changed. During Soviet times, the mation, also provide evidence that vation). This decline in income and major threat to wolves was the large- wolf populations have actually de- employment coincided with the dis- scale government sponsored wolf clined over the last decade. mantling of much of the social safety control programs. When the net that was in place during the Soviet economy experienced a recession, Discussion times. Medical care and education are there was less money to spend on There are two questions that need no longer free and subsidies for food these programs and wolf populations to be addressed. One is why, dur- and utilities have either been eliminated were able to rebound (Bibikov 1993). ing the current economic hard or sharply reduced (Anderson and Currently, the major threat is a lack times, are wolf populations declin- Pomfret 2000). This has resulted in of natural resources. The prey base ing rather than increasing, as they close to 80% of the population of for wolves has declined significantly have in the past during Soviet Kyrgyzstan living in poverty (Minis- in less than 10 years due to habitat de- times? The other is why the current try of Environmental Protection 1998). struction, increased human exploitation perception is that wolf populations The increase in poverty and un- of game species, and a decline in do- are increasing when populations employment and the changes in gov- mestic livestock. In addition, while may actually be declining? In order ernment subsidies for social pro- there are currently no government em- to answer these questions, we must grams are only part of the story. In ployees being paid to hunt wolves, understand how the economy and addition, there have been substantial there are still bounties in place that en- lifestyle of the people have changed changes in the structure of the agri- courage private hunting of wolves.

Vol. 18 No. 4 2001 Endangered Specles UPDATE 145 populations are stable or increasing appears to be common in these re- gions. This perception in Kyrgyzstan, coupled with the in- creased economic impact of wolf depredation due to the increase in poverty and private ownership of livestock, appears to be exerting in- creasing pressure on the government to re-institute wolf control programs (Kurmanaliev, personal cornrnunica- tion). This could be devastating if wolf populations are already in decline. The Kyrgyz Republic is only one small part of the former Soviet Union, but most former republics are experiencing similar social and eco- Over 90% of the countryside of Kyrgystan is mountainous (photograph by nomic problems. We need to more CJ Hazell). fully investigate how these changes are affecting wildlife communities. Most of this region has reported that The reason that people perceive true. The increased economic hard- wolf populations are stable. We wolf populations as increasing rather ship for farmers resulting from wolf should be skeptical of this. than decreasing may be due to the depredation is coupled with a decline changes in both the economy and in in wild prey species. The wolves Literature cited the lifestyle of people. Economically, have fewer alternatives to livestock. Anderson, K. and R. Pornfret. 2000. Living standards during transition to a market before independence, individuals This may be resulting in wolves com- economy: the Kyrgyz Republic in 1993 and didn't experience significant changes ing closer to villages as well as a pos- 1996. Journal of Comparative Economics in living conditions during a reces- sible increase in depredation. The lat- 28502-523. sion because of the regulated, cen- ter is difficult to substantiate due to Bibikov, D.I. 1993. The wolf in the cornrnu- the lack of documentation of depre- nity of independent states (former Soviet tralized economy that existed during Union). Pp 22-24 In Promberger, C. and W. the Soviet times. The social safety dation rates. These conditions could Schroeder, eds. Wolves in Europe: Status net that existed guaranteed access to easily result in the perception that and Perspectives. Proceedings of a Work- employment and basic amenities. In wolves are increasing in number sim- shop in Oberarnmergau, Germany April 2- addition, most livestock was gathered ply because their visibility and eco- 5, 1992. Munich Wildlife Society. Ministry of Environmental Protection. 1998. into large state-owned herds. Wolf nomic impact may be increasing. Kyrgyz Republic biodiversity strategy and ac- depredation resulted in the taking of tion plan. MEP. Bishkek, Kyrgyz Republic. a few head of sheep from very large Conclusion Osh Oblast Hunting Institution. 1988-1999. Re- flocks so proportionately the losses The former Soviet Union comprised ports on wildlife populations. Osh, a very large geographic area. Since Kyrgyzstan. were not as significant. Osh Oblast Subsistence Farm Administration. Since independence, the state- its breakup, the former republics are 1987-1997. Total Counts of Livestock and owned herds have been broken up all experiencing profound economic Domestic Birds in AU Categories. Osh Oblast and distributed to individuals, result- and social upheaval. These changes Committee on Subsistence. National State ing in many small, privately owned are not only affecting people, but are Committee on Statistics. Kyrgyz Republic. Route, B. and L. Aylsworth. 1999.1999 World herds. In addition, the economic also having a significant impact on Wolf Status Report. International Wolf 9(3) downturn has resulted in a significant wildlife. Most of the resulting de- increase in poverty and a drop in the clines in wildlife populations are be- Theberge, J. B. 199 1. Ecological classification, standard of living for most people. ing acknowledged. Wolves, how- status, and management of the gray wolf, Ca- ever, have not been recognized as a nis lupus, in Canada. The Canadian Field- When sheep were taken from a state- Naturalist 105(4):459-463 owned herd, it didn't result in indi- species that may also be experienc- World Bank. 1999. Kyrgyz Republic: update vidual farm workers experiencing ing negative impacts from the current on poverty in the Kyrgyz Republic. Report economic hardship. This is no longer situation. The perception that wolf 19425-KG. World Bank.Washington. D.C.

146 Endangered Species UPDATE Vol. 18 No. 4 2001 Gray Wolf Restoration in the Northwestern United States Ed Bangs Wolf Recovery Coordinator, U.S. Fish and Wildlife Service, 100 N. Park, #320, Helena, MT 59601; (406) 449-5225 x204; (fax) (406) 449-5339; [email protected]; http://midwest.fws.gov/wolf;http://mountain-prairie.fws.gov/wolf/annualrptOO/ Joe Fontaine Assistant Wolf Recovery Coordinator, U.S. Fish and Wildlife Service, 100 N. Park, #320, Helena, MT 59601; (406) 449- 5225 x206; joe-fontaineQfws.gov Mike Jimenez Project Leader Wyoming Wolf Recovery, U.S. Fish and Wildlife Service, 190 N. 1st St., Lander, WY 82520; (307) 332- 7789; [email protected] Tom Meier Wolf Biologist, U.S. Fish and Wildlife Service, c/o MT Fish, Wildlife and Parks, 490 N. Meridian Rd., Kalispell, MT 59901; (406) 751-4581; [email protected] Carter Niemeyer Project leader for ldaho Wolf Recovery, U.S. Fish and Wildlife Service, 1387 S. Vinnel Way, Rm 368, Boise, ID 83709; (208) 378-5639; [email protected] Doug Smith Project Leader, Yellowstone Wolf Recovery, Yellowstone Center for Resources, P.O. Box 168, Yellowstone National Park, WY 82190; (307) 344-2242; [email protected] Kerry Murphy Biologist, Yellowstone Wolf Recovery, Yellowstone Center for Resources, P.O. Box 168, Yellowstone National Park, WY 821 90; (307) 344-2242; kerry-murphy @nps.gov Deb Guernsey Program Assistant, Yellowstone Wolf Recovery, Yellowstone Center for Resources, P.O. Box 168, Yellowstone National Park, WY 821 90; (307) 344-2242; deb-guernsey @nps.gov Larry Handegard Montana State Director, Wildlife Services, USDAIAPHIS, P.O. Box 1938, Billings, MT 59103; (406) 657-6464; larry.l.handegardQaphis.usda.gov Mark Collinge ldaho State Director, Wildlife Services, 9134 W. Blackeagle Dr., Boise, ID 83709; (208) 378-5077, mark.d.collingeQaphis.usda.gov Rod Krischke Wyoming State Director, Wildlife Services, P.O. Box 59, Casper, WY 82602; (307) 261-5336; rod.f.krischkeOaphis.usda.gov John Shivik Leader Predation Management Project, WS National Wildlife Research Center, 4101 LaPorte Ave, Fort Collins, CO 80521; (970) 266-6088; john.shivikQaphis.usda.gov Curt Mack Project Leader Nez Perce Tribe Wolf Recovery, 1000 Mission, McCall, ID 83638; (208) 634-1061; cmackOnezperce.org Issac Babcock Nez Perce Wolf Biologist, 1000 Mission, McCall, ID 83638; (208) 634-1061; ibabcock.hotbot.com Val Asher Wolf Biologist, Turner Endangered Species Fund, 1123 Research Dr., Bozeman, MT 59718; (406) 556-8514; valasherd montana.net Dominic Domenici Wyoming Senior Resident Agent, U.S. Fish and Wildlife Service, 100 East B St., Federal Bldg Rm 41 30, Casper, WY 82602; (307) 261-6365; [email protected]

Abstract Gray wolf (Canis lupus) populations were eliminated from Montana, Idaho, and Wyoming, as well as adjacent southwestern Canada by the 1930s. After human-caused mortality of wolves in southwestern Canada began to be regulated in the 1960s)populations began expanding south- ward. Dispersing individuals occasionally reached the northern Rocky Mountains of the United States, but lacked legal protection there until 1974, after passage of the Endangered Species Act of 1973. In 1986, wolves from Canada successfully raised a litter of pups in Glacier National Park, Montana, and a small population was soon established. In 1995 and 1996, wolves from

Vol. 18 No. 4 2001 Endangered Species UPDATE 147 western Canada were reintroduced to remote public lands in central Idaho and Yellowstone National Park. These wolves were designated as nonessential experimental populations to increase managementflexibility and address local and state concerns. Wolfrestoration is rapidly occurring in Montana, Idaho, and Woming, and there were at least 28 breeding pairs in Decem- ber 2000. There are now about 63 adult wolves in northwestern Montana, 192 in central Idaho, and 177 in the Greater Yellowstone area. Dispersal of wolves between Canada, Montana, Idaho, and Wyoming has been documented. Occasional lone wolves may disperse into adjacent states, but population establishment outside of Montana, Idaho, and Wyoming is probably not imminent. The gray wolfpopulation in the northwestern U.S. should be recovered and, depending on the completion of state and tribal wolf conservation plans, could be proposed to be removed from Act protection within three years. Wolf restoration has proceeded more quickly and with more benefits, such as public viewing than predicted. Problems, including confirmed livestock depre- dations, have been lower than estimated. The Service led interagency recovery program focuses its efforts on achieving wolf recovery while addressing the concerns of people who live near wolves. Wolves have restored an important ecological process to several large wild areas in the northern Rocky Mountains of the U.S. The program has been widely publicized and is generally viewed as highly successful.

Wolves in northwestern control actions in response to high report was completed in May 1991, Montana livestock depredations and subse- but Congress chose not to act on the Sixty years after being nearly exter- quent poor pup production (Bangs et Committee's recommendation, which minated from the lower 48 states, the al. 1998). Wolf numbers have only included wolf reintroduction and gray wolf (Canis lupus) was listed slightly increased since 1997. In more flexible wolf management than under the Endangered Species Act 2000, there were an estimated 63 was normally allowed under the (ESA) in 1974 and was eventually re- wolves in about a dozen groups, but ESA. All these reports, and all sub- stored to Montana, Idaho, and Wyo- only six of those successfully repro- sequent investigations, made it clear ming. Wolves were once common duced. Most wolves in NW Montana that reintroducing wolves in throughout North America but were live in a mix of private and public Yellowstone National Park and cen- deliberately exterminated in the land west of the Continental Divide. tral Idaho was feasible and would lower 48 states (except northeastern ultimately result in wolves attempt- Minnesota). Wolves remained abun- Wolf reintroduction in ing to recolonize areas throughout dant in much of Canada and Alaska. Yellowstone National Park Montana, Idaho, and Wyoming and Recovery began in northwestern and central Idaho far outside the reintroduction areas. (NW) Montana in the late 1970s by In 1988 and 1990, Congress directed In late 1991, Congress directed natural dispersal from nearby ex- the National Park Service to prepare the Fish and Wildlife Service (FWS) panding Canadian wolf populations a series of reports on the potential to lead preparation of an environmen- (Pletscher et al. 1997). Wolves first effects of reintroducing wolves to tal impact statement (EIS) to exam- denned in NW Montana in Glacier Yellowstone National Park (YNP ine the effect of reintroducing wolves National Park in 1986 (Ream et al. 1990). Wolf depredation on live- to Yellowstone National Park and 1989). Wolf numbers steadily in- stock, wolf predation on wildlife, central Idaho (FWS 1994). The plan- creased until 1996, when there were land-use restrictions, tourism, other ning and public involvement effort a minimum of 70 wolves in seven predators including grizzly bears took two years to complete. By the different packs that lived solely in (Ursus arctos), diseases, and a wide time it was finished the Service had NW Montana. An unusually severe variety of other issues were evalu- distributed over 750,000 documents, winter in 1996-97 caused a 30 to 50% ated. In 1990, Congress established conducted over 130 public meetings decline in the white-tailed deer a Wolf Management Committee, con- and hearings, and reviewed 170,000 (Odocoileus virginanus) populations, sisting of federal, state, and private public comments. The decision was the primary prey of those wolves. special interest groups to try to forge to reintroduce wolves to both The number of wolves dropped to a political compromise on the issue Yellowstone and central Idaho as just over 50 in five packs in 1997, of wolf reintroduction in both nonessential experimental popula- largely as a result of agency wolf Yellowstone and central Idaho. Their tions, the most flexible classification

148 Endangered Species UPDATE Vol. 18 No. 4 2001 for species listed under the ESA. The area was worth to them). Similar pre- primarily managed by the Nez Perce decision was approved in spring 1994 dictions were made for the central Tribe and Wildlife Services, under a by both the Secretary of the Interior Idaho area. Depending upon their cooperative agreement with the FWS. (FWS, National Park Service and distribution, more than 100 adult- Reintroduced wolves adapted Bureau of Land Management) and sized wolves would proportionally better than predicted and only two the Secretary of Agriculture (Wild- increase impacts above those pre- years of reintroduction were required life Services and Forest Service). dicted in the EIS. To date, at least rather than the three to five years that The EIS predicted that a recov- the trends in these predictions appear were predicted (Fritts et al. 1997). In ered wolf population (a minimum of to have been fairly accurate. It will December 2000, the population esti- 10 breeding pairs, estimated to be take time before wolf numbers and mate was 177 wolves in 13 breeding about 100 adult-sized wolves) in the distribution stabilize and the true ef- groups in the Yellowstone area and Yellowstone area would kill an aver- fect of having wolves back in these 192 wolves in 9 breeding groups in age of 19 cattle (Bos sp.), 68 sheep areas can be ascertained. Idaho. To date, wolves have settled (Ovis aries), and up to 1,200 ungu- The restoration of wolves to pub- primarily on remote public lands, but lates (primarily elk) annually. This lic lands in the western United States, that will change as the population would not affect hunter harvest of particularly Yellowstone National expands and more wolves disperse male ungulates, but could reduce Park, was proposed as early as the beyond where wolf packs currently hunter harvest of female elk (Cewus 1940s. After years of direct involve- exist. Dispersing wolves will in- elaphus), deer (Odocoileus sp.), and ment by Congress and exhaustive creasingly try to occupy private lands moose (Alces alces) in some herds. public involvement and planning, 35 used for livestock production; this Hunter harvests or populations of wolves were reintroduced via hard will increase the rate of livestock dep- bighorn sheep (Ovis Canadensis), (immediate) release to wilderness ar- redations and agency control. Except mountain goats (Oreamnos eas in central Idaho, and 3 1 were soft for a few temporary closures to pro- americanus), or antelope (Antilopra released in Yellowstone National tect wolf viewing opportunities americana) would not be affected. Park, Wyoming in January 1995 and around active dens in Yellowstone Bison (Bison bison) would not be January 1996 (Fritts et al. 1997; National Park, and restricting some preferred prey. Wolf predation may Bangs et al. 1998). Those wolves, M-44 use, the wolf restoration pro- reduce populations of elk five to originally from Canada, were desig- gram has caused no land-use restric- 30%, deer three to 19%, moose seven nated as nonessential experimental tions that might disrupt traditional percent, and bison up to 15%. The populations to increase management human activities such as logging, presence of wolves would not change flexibility over what is normally al- mining, livestock grazing, hunting, uses of public or private land except lowable for species listed under the trapping, or wildland recreation. Over for potential use of M-44 cyanide ESA. Examples of this flexibility 70,000 visitors toYellowstone National devices, used to control coyote (Ca- are: landowners could harass wolves Park have seen wolves and public in- nis latrans) damage, in areas occu- at any time; livestock producers terest in them is extremely high. pied by wolves. Visitor use was pre- could shoot wolves seen attacking dicted to increase five to 10%. At livestock; wolves could be relocated Wolf research wolf recovery, annual economic if they significantly impacted wild Between 1979 and the late 1990s, losses were estimated to be $1 87,000 ungulate herds (as defined in ap- extensive research on wolves in NW to $465,000 in hunter benefits (what proved state wolf management Montana was supported by a host of hunters said hunting female elk was plans); there would be virtually no state and federal resource manage- worth to them), $207,000 to land-use restrictions; the Service. ment agencies. Field work and data $4 14,000 in potential reduced hunter could use special permits to take analysis were carried out largely by expenditures (what hunters of female wolves for various management rea- graduate students and the University elk said they would have spent hunt- sons; and funding was offered for of Montana. Those studies investi- ing), and $1,888 to $30,470 in poten- state and tribal leadership in wolf re- gated the relationships between tial livestock losses. Annual in- covery actions (Bangs and Fritts wolves and other wildlife, including creased visitor expenditures were es- 1996). Currently wolves in Wyo- white-tailed deer, elk, and moose, timated at $23 million and the exist- ming and Montana are primarily other predators such as mountain li- ence value of wolves was estimated managed by the FWS, National Park ons (Felis concolor) and coyotes, and at $8.3 million (what people believed Service (in Parks), and USDA Wild- livestock (Kunkel and Pletscher having wolves in the Yellowstone life Services. In Idaho, wolves are 1999; Kunkel et al. 1999; Kunkel

Vol. 18 No. 4 2001 Endangered Species UPDATE 149 1997). This research indicated ing dogs, were apparently killed as calf elk killed by wolves. Bison are wolves were just another predator on competitors rather than prey. Wolves difficult to kill and few wolves have wild ungulates, neither much less nor commonly fed on carrion of both learned to do so effectively (Smith much more effective than other na- livestock (carcass dumps) and wild et al. 2000). Somewhat surpris- tive predators, such as mountain li- ungulates (road and train kills, ingly, to date no Bighorn Sheep ons, black (Ursus americanus) and unretrieved hunter-killed game, and have been confirmed killed by grizzly bears, or coyotes. Wild preda- gut piles). In some instances, abun- wolves in either area. tors, including wolves, typically dance of natural prey and relative Carcasses of elk killed by wolves killed more of the most vulnerable of vulnerability of livestock affected were utilized by a wide variety of ungulates (injured, sick, or very how often wolves attempted to attack other wildlife species and provided a young and very old individuals) than livestock. Sick or wounded livestock year-long food source that would did human hunters. Wolf predation or small livestock, such as calves or likely increase overall wildlife diver- in combination with other factors sheep, appeared particularly vulner- sity. Coyote numbers in some areas such as winter weather, human hunt- able to wolf predation. But often, may have been reduced by half be- ing, other predators, and habitat con- wolves appeared to attack livestock cause of wolves killing coyotes. ditions, contributed to a decline in without any predisposing factors and Mountain lions and wolves tend to white-tailed deer and elk in the North nearly all wolf packs with regular kill the same types of prey, but lions Fork of the Flathead River. Moose exposure to livestock sporadically are usually confined to more rugged populations apparently were not as caused depredations. steep and vegetated terrain, while affected by these same circum- A large number of studies and wolves preferred flatter terrain and stances. As a result of that prey de- research are currently being con- made more use of open habitat. Griz- cline, wolf numbers in that area dra- ducted on wolves in the Yellowstone zly bears often usurped wolf killed matically declined, from nearly 30 and central Idaho experimental areas ungulates. Studies are investigating wolves in three packs during the most so that accurate information can be the effect of wolves on elk distribu- intensive research in the early 1990s used to better manage wolf popula- tion on winter feeding grounds in to a few individuals that did not pro- tions and expand the level of knowl- Wyoming, but tentative results sug- duce pups in 1999 or 2000. Wolves edge about wolves. Wolf predation gest little effect other than elk appear often trailed mountain lions to take studies indicated elk were more than to be more wary and may prefer over their kills and killed a few li- 90% of the prey killed by wolves in larger groups and more open habitat ons. Direct competition for the types Yellowstone. Kill rates were about when wolves are present. Earlier of ungulates that are most vulnerable 15 elk per wolf per year. In Idaho, studies in Montana indicated that to predation was likely the main im- wolves also preyed mainly on elk, but wolves did not change ungulate dis- pact that wolves would have on li- wolves there killed a higher propor- tribution on natural winter ranges, ons. Wolves also killed a few coy- tion of mule deer (Odocoileus but apparently caused ungulates to otes. While wolves displaced lions hemionus). Wolf kills were more be more wary and to temporarily and coyotes from ungulate carcasses, likely to be in open habitats and the retreat to thicker cover when wolf kills were often usurped by griz- remains scattered, compared to wolves were present. zly bears. Studies of wolf genetics mountain lion kills that were often A recent study funded and initi- and dispersal indicated that genetic covered and hidden in thick cover. ated by the Nez Perce Tribe and a host diversity was high and likely not a This gave a visual impression that of federal agencies and local live- management concern, as long as op- wolves killed more deer and elk than stock producers found that confirmed portunity for occasional dispersal mountain lions, but a lion actually livestock losses may be a fraction of from wolf populations in Canada and kills more ungulates per year than actual losses under some circum- other U.S. recovery areas in Idaho does a wolf. Annual wolf kill rates stances. That study determined the and Wyoming was maintained (Boyd typically average about 20 adult deer cause of death and detection rate of and Pletscher 1999). or 12 adult female elk per year, while 220 radio-tagged livestock calves of Research indicated that although adult lion kill rates can be twice as about 700 on large, very remote, and wolves often lived near livestock high. Both wolves and lions tended heavily forested USDA Forest Ser- (primarily cattle) and other domestic to prey on the most vulnerable wild vice grazing allotments. After two animals, conflicts were uncommon. ungulates such as calves and very old years, pneumonia killed the most Dogs, almost exclusively hunting females. Calf elk killed by lions in marked calves, but wolf predation hounds and livestock guard and herd- Idaho were in better condition than was the second leading cause of

150 Endangered Species UPDATE Vol. 18 No. 4 2001 death. Sample sizes were very small, wolf depredation and control actions vate property, to shoot wolves on- but as many as 5.7 calves may have still result in major local news cov- sight. We have allowed landowners died from wolf predation for every erage. To the general public, this to non-injuriously harass wolves at one discovered by normal livestock probably greatly exaggerates both the any time. We have trained and then herding practices. Wolves killed role of wolves as livestock predators issued cracker shells and less-than- calves that were the lowest weight, and the level of agency control. Since lethal munitions (12-gauge bean- least guarded by people, nearest to 1987, livestock producers who expe- bag or rubber bullet shells) to pri- an active wolf den, and in the heavi- rienced confirmed or highly probable vate landowners so they could in- est forest cover, suggesting that wolf-caused losses in Montana, juriously harass any wolves near wolves tested and hunted cattle like Idaho, and Wyoming have been their livestock or property. wild prey and attacked the most vul- compensated about $155,000 by a nerable animals. private compensation fund admin- Litigation istered by the Defenders of Wild- Several lawsuits were filed over the Livestock depredations life, who support wolf recovery and reintroduction program, by a wide Since 1987, annual confirmed mini- management efforts. variety of groups, including the Si- mum livestock losses in NW Mon- erra Club Legal Defense Fund who tana totaled 82 cattle, 68 sheep and Minimizing livestock conflicts supported and the American Farm seven dogs. As a result, 41 wolves The Service is evaluating a wide va- Bureau Federation who opposed wolf were killed and 32 were moved. Dep- riety of alternative methods to pre- restoration. The lawsuits were redations averaged 5.8 cattle, 4.8 vent or reduce conflicts with live- pooled into a single case that ques- sheep, and less than one dog annu- stock in addition to relocating or kill- tioned whether the Service's use of ally. Agency control killed an aver- ing problem wolves. The experimen- an experimental population designa- age of three wolves per year. On av- tal population rules and the recently tion for reintroduced wolves illegally erage, less than six percent of the proposed. special rule for wolves reduced protection of wolves that wolf population is annually affected listed as threatened would allow for might naturally wander into the ex- by agency wolf control actions harassment and killing of problem perimental areas. To date, no natu- (Bangs et al. 1995). Minimum con- wolves. In cooperation with USDA rally dispersing wolves have been firmed livestock losses have annually Wildlife Services and private conser- found in the Yellowstone area, but at averaged about 3.6 cattle, 27.8 sheep, vation organizations we have: used least three wolves from NW Montana and 3.8 dogs in the Yellow stone area, light and siren devices, including have dispersed into the central Idaho and 9.2 cattle, 29.4 sheep, and 1.8 models triggered by the signals from area. The Wyoming District Court dogs in central Idaho. In addition, individual radio-collared wolves; es- eventually ruled against the Service's one newborn horse (Equus sp.) was tablished barriers to wolves using position in December 1998 and or- killed in the Yellowstone area. In guard animals, flagging and fencing; dered all the reintroduced wolves re- total there have been 146 cattle, 356 provided extra surveillance of live- moved, but stayed its own decision sheep and 35 dogs confirmed killed stock with herders or agency person- pending appeal. That case was then by wolves from 1987 until January nel; harassed and moved andlor pro- reviewed by the Tenth Circuit Court 2001. Since 1987, the Service and vided supplemental food to wolves of Appeals in Denver, Colorado. USDA Wildlife Services have killed that established dens and rendezvous Their ruling in January 2000 over- 41 wolves in NW Montana, 18 in sites in livestock grazing pastures; turned the Wyoming lower court rul- central Idaho, and 26 in the initiated research using electronic ing. The Tenth Circuit endorsed and Yellowstone area because of conflicts dog training collars to teach wolves validated the legality of the Service's with livestock. The rate of confirmed not to attack livestock; provided live- authority and the wolf reintroduction wolf-caused livestock losses and the stock producers radio telemetry re- program. None of the losing parties number of wolves that have been re- ceivers so they could closely moni- appealed to the Supreme Court per- moved in agency control actions is tor wolves near their livestock; and haps because several months earlier, one-third to one-half of the levels helped provide alternative pasture to in a closely related case that involved predicted in the EIS. Despite lower reduce livestock and wolf encounters. the illegal killing of an reintroduced than expected losses and less wolf We have permitted livestock produc- wolf, the Ninth Circuit Court of Ap- control than predicted, wolf depreda- ers to shoot wolves actually seen at- peals in California had also ruled tions and control remain inordinately tacking livestock, and in a few strongly in favor of the Service's au- controversial. Even the most routine chronic cases of depredation on pri- thority and the Supreme Court had

Vol. 18 No. 4 2001 Endangered Species UPDATE 151 refused to hear an appeal. The only delisting proposal, including assur- vation of wolves in a changing world. Ca- unresolved litigation involves a Wyo- ance that state wolf management nadian Institute$ Alberta. ming rancher who suffered several plans would conserve wolves above Boyd, D.K, and D.H. Pletscher 1999, confirmed wolf-caused livestock and recovery levels, the affected states Charcteristics of dispersal in a colonizing pet depredations, and suspects he had and tribes could manage wolves with- wolf population in the central ~ocky many other unconfirmed losses. He out federal oversight, except for the Mountains. Journal of Wildlife Manage- claims that the government reintro- five-year post recovery monitoring ment 63:1094-1108. Fritts, S.H., E.E. Bangs, J.A. Fontaine, M.R. duction program resulted in an un- period that is required by the ESA. Johnson, M,K, Phillips, E.D. Koch, and compensated "taking" of his private State and tribal management pro- J.R. Gunson. 1997. Planning and imple- property and lifestyle. Other litiga- grams will likely allow wolves to be menting a reintroduction of wolves to tion on a wide variety of wolf man- killed in defense of life and property Park and Idaho. Restoration Ecology 57-27, agement issues is almost certain be- and in regulated public harvest pro- Fritts, S.H. and L,N, Carbyn. 1995 Popu- cause of the strong symbolism of grams, just as other large predators lation viability, nature reserves, and the wolves to various special interest in these states are managed. Ulti- outlook for gray wolf conservation in groups and the public, both at the lo- mately, wolf numbers (above mini- N~*hAmerica. Restoration Ecology 3:26- cal and national level. mum 38. levels) and pack Kunkel, K.E. 1997. Predation by wolves and The Service-led interagency wolf distribution will be determined by other large carnivores in northwestern recovery program focuses its efforts state and tribal wildlife management Montana and southeastern British Colum- on achieving the wolf recovery goal agencies. Once recovery goals have bia. Ph.D. dissertation, University of while addressing the concerns of been achieved, delisting and a return Montana, MissOula$MT. Kunkel, K.E., T.K. Ruth, D.H. Pletscher, and people who live near wolves. Over to sole state and tribal management M.G, Hornocker 1999. Winter prey se- 85% of all known wolf mortalities are will signal the final success of the lection by wolves and cougars in and near caused by people, and the majority ESA at recovering the once imper- Glacier National Park, Montana. Journal of those are a result of agency wolf iled gray wolf in the northern Rocky Management63:901-910. control actions (Bangs et al. 1998). Mountains of the U.S. Kunkel, K.E. and D.H. Pletscher. 1999. Spe- cies-specific population dynamics of The key to successfully completing cervids in a multipredator ecosystem. Jour- wolf restoration efforts will depend Acknowledgments nal of Wildlife Management 63:1082- on maintaining some connectivity Literally hundreds of people have 1093. between the few remaining areas of contributed in a multitude of ways to 'letscher* D*H., R*R. Ream$ D.K. M.W. Fairchild, and K.E. Kunkel. 1997. large wild habitat remaining in the wolf restoration. It is impossible for Population dynamics of a recolonizing western U.S. and tolerance of wolves us to individually recognize and population in the ~~~k~ Mountains, by the local rural residents (Fritts and thank all of them in this paper, but . Journal of Wildlife Management 61:459- Carbyn 1995). without strong interagency coopera- 465 tion and public involvement and tol- Ream, R.R., W. Fairchild, D. K. Boyd and A. Blalresley. 1989. Firs wolf den in Wolf recovery erance, the successes so evident to- western United States in recent history. Wolf populations should be fully re- day would not have been possible. Northwestern Naturalist 70:39-40. covered (30 breeding pairs with eq- Smith, D.S., L.D. Mech, M. Meagher, W.E. uitable distribution throughout the Literature cited Clark, R. Jaffe, M.K. Phillips, and J.A. Mack. 2000. Wolf-Bison interactions in Bangs, E.E. and S.H. Fritts. 1996. Reintro- three recovery areas for three succes- Yellowstone National Park. Journal of sive years) and will no longer need ducing the gray to Idaho and Mammalogy 81:1128-1135. Yellowstone National Park. Wildlife So- U.S. Fish and Wildlife Service. 1994. The protection under the ESA by 2003. ciety Bulletin 24:402-4i3. reintroduction of gray wolves to As a result of dispersal by wolves Bangs E.E., S.H. Fritts, J.A. Fontaine, D.W. Yellowstone National Park and Central from Canada and the combination of Smith, K.M. Murphy, C.M. Mack, and Idaho. Final Environmental Impact State- C.C. Niemeyer. 1998. Status of gray wolf reintroduction from two areas in ment. USFWS, Helena, MT. restoration in Montana, Idaho, and Wyo- Canada, genetics not be a _ng, wildlife society ~~ll~~i~26:785- Yellowstone National Park, U.S. Fish and Wildlife Service, University of Wyoming, tor in wolf population viability as 798. University of Idaho, Interagency Grizzly long as some connectivity is main- Bangs, E.E., S.H. Fritts, D.A. H~S,J.A. Bear Study Team, and University of Min- Fontaine, M.D. Jimenez, W.G. Brewster, tained (Boyd and Pletscher 1999). nesota Cooperative Parks Studies Unit, and C.C. Niemeyer. 1995. Control of en- oncethe recovery goal is achieved, editors. 1990. Wolves for Yellowstone? dangered gray wolves in Montana. Pp A report to the United States Congress. Vol be proposed to be 127-134 in L.N. Carbyn, S.H. Fritts, and I. Research and analysis. Mammoth, Wyo. delisted. After extensive public and D.R. seip, editors. ~~~l~~~ and censer- professional review of the Service's 586 pp.

152 Endangered Species UPDATE Vol. 18 No. 4 2001 The Feasibility of Gray Wolf Reintroduction to the Grand Canyon Ecoregion Paul G. Sneed Environmental Studies , Master of Arts Program, Prescott College, 4906 Box Canyon Road, Billings, MT 59101;(406) 245-9117; (fax) (406)245-0787; [email protected]

Abstract As part of a regional conservation planning initiative, this study is being undertaken to determine the biophysical and socioeconomic feasibility of reestablishing a top carnivore, the gray wolf (Canis lupus), in the Grand Canyon Ecoregion (GCE). The GCE is a roughly 1.5 million km2 area located on the southern Colorado Plateau. The last remaining gray wolves were probably eradicated in the 1920s and 1930s. Because of an interest in restoring extirpated native species to this ecoregion, and the desire to increase the size of the gray wolfmetapopulation in the Southwest, there is need for an objective and spatially explicit landscape-scale model of potential gray wolfhabitat. Thefirst phase of this conservation GIS analysis involves utilizing six habitat characteristics or factors-vegetation cover, suface water availability, prey density, human population density, road density, and land ownership-to identify and describe potential reintro- duction sites in the Arizona section of the Grand Canyon Ecoregion. Initial results show that there are at least two localities in northern Arizona suitable for reintroduction of around 100 wolves. This paper is a preliminary report on observations, results, and some recommendations deriving from the feasibility study.

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Introduction Crumbo 1998). This paper, how- Regardless of inaccuracies in the Conservation biologists have shown ever, will focus on current, prelimi- historical record, a partial picture of that large or top carnivores are often nary results from research done on where wolves occurred prior to their keystone species whose removal a limited number of factors in the extermination in the Southwest can jeopardizes the maintenance of eco- northern Arizona section of the still be pieced together. These records logical integrity in large-scale ecosys- ecoregion (see Figure 1). show that at least small populations tems (Soule and Noss 1998;Terborgh of wolves were found throughout the et al. 1999). Therefore, conservation Historic occurrence and woodlands and forests of northern planners interested in restoring and taxonomic position Arizona (Brown 1984). For example, protecting large ecosystems or To accurately reconstruct the his- from these records we know that there ecoregions emphasize recovery of top toric distribution of gray wolves in were at least 30 wolves on or near the predators. The primary goal of this the GCE is challenging for a vari- North Kaibab because of the number study-which is supported by the ety of reasons. Nineteenth century reported killed between 1907 and Grand Canyon Wildlands Council, writers often accidentally or pur- 1926 (Russo 1964). Brown (1984) Defenders of Wildlife, and Prescott posefully misidentified coyotes claimed that "the last wolf in this part College-is to determine the capabil- (Canis latrans), wolves, and wolf- of northern Arizona was taken on the ity and suitability (together, the fea- dog hybrids (Gipson et al. 1998). Paria Plateau about 1928", but a sibility) of reintroducing one top car- Wolf hunters and trappers some- former Civilian Conservation Corps nivore, the gray wolf (Canis lupus), times exaggerated the number of worker recently reported that he saw to the Grand Canyon Ecoregion wolves in an area to enhance their wolves on three different occasions (GCE) (Figure 1). job security and occasionally mis- in 1935 on the North Rim of Grand Ultimately, this study will ad- represented where a wolf was killed Canyon National Park (GCNP) dress 26 factors or aspects, grouped in order to claim a local bounty. (Leslie, personal communication). into two dimensions-biophysical Furthermore, the widespread use of Moreover, "as recently as March 3, and human-that are expected to af- poisons meant that many animals, 1948, assistant chief ranger A.L. fect the feasibility of wolf recovery including wolves, were dispatched Brown reported wolf tracks in fresh in the entire GCE (Sneed and without any record of their death. snow in the area of Bright Angel

Vol. 18 No. 4 2001 Endangered Species UPDATE 153 to adequate food supplies, security from human disturbance and perse- cution are important factors affect- ing the suitability of a landscape for wolf recovery. At this stage in the research, three critical human di- mension aspects are considered: human population density, road density, and land status.

Biophysical factors Several reintroduction studies (e.g., Mladenoff et al. 1995) suggest that gray wolves, at least those living Figure 1. Grand Canyon ecoregion dispersed into the southeast- south of the Arctic, tend to prefer for- em part of the GCE. ested landscapes. Historically, in the Southwest, wolves were most com- Point" on the North Rim of GCNP Habitat capability and monly found associated with wood- (Hoffmeister 1971). Finally, the last suitability mapping lands and montane forests (Groebner wolf inhabiting the Mogollon Rim Restoration of viable large carnivore et al. 1995; FWS 1996). When ob- area in the southern part of the GCE populations is probably among served elsewhere, such as in grass- was reportedly taken in 1942 society's greatest challenges, requir- lands, they were probably simply (Hoffmeister 1986). Clearly, gray ing extraordinary innovation and co- passing through as they moved be- wolves occurred within the Grand operative management on an tween their preferred habitat of for- Canyon Ecoregion well into the twen- ecoregional scale (Paquet and Hack- ested highlands. Figure 2 maps the tieth century, although their exact man 1995). Furthermore, solutions distribution of these two vegetation numbers and range will probably to large predator conservation are types, as well as others such as never be known with certainty. economic, sociological and political shrublands and grasslands. This fig- Due to the taxonomic splitting (human dimension issues), as well as ure plainly illustrates a broad band of approach of the time, Young and biological and ecological (biophysi- forestlands-woodlands extending Goldman (1944) identified 23 sub- cal factors) (Clark et al. 1996). The north-south from the Kaibab Pla- species of North American gray feasibility of wolf recovery depends teau, through the Flagstaff area to wolves (based on skull measure- on the capability and suitability of the Mogollon Rim, interrupted only ments, pelage color, and size) and habitat for sustaining wolf popula- by the Grand Canyon and urbanized mapped their geographic distribution. tions. Although many factors can and areas such as Flagstaff. Other ar- Two of these 23 nominal subspe- should be considered, the ultimate de- eas of woodland/forest vegetation cies-C. I. mogollonensis (the Ari- terminants are ungulate prey and hu- types are found in isolated moun- zona wolf), C.l. youngii (the Great man impact (Fuller et al. 1992) or, put tain areas of the Arizona Strip as Basin or Intermountain wolf), and, another way, sustenance and security. well as the Hualapai and Navajo possibly, C.I. baileyii (the Mexican Course screen landscape-scale Indian Reservations. wolf)-inhabited the Grand Can- habitat mapping for the Arizona por- Because wolves require large yon Ecoregion (Brown 1984; FWS tion of the GCE (see Figure 1) has amounts of water to aid digestion 1996). Development of similar been done following other .similar (Lopez 1978; Mech 1970), several classification schemes continued studies (Mladenoff et al. 1995; studies of wolves in the Southwest into the 1970s (e.g., Hall and Quinby et al. 1999; Ratti et al. 1999; (Groebner et al. 1995; FWS 1996) Kelson 1959) until some taxono- Wydeven et al. 1998). Various bio- and elsewhere (Quinby et al. 1999) mists began questioning the split- physical factors can be considered in have suggested that the availability ting tradition of wolf . evaluating the capability of habitat to of free water is an important deter- Modern lumping systems of wolf support wolves, but this study focuses minant of gray wolf abundance and taxonomy are based on multivariate on vegetation cover, surface water distribution. Figure 3 illustrates the statistical analysis of large sample availability, and, most importantly, distribution of currently mapped sizes and confirmed by the results of ungulate prey abundance. In addition lakes, springs, and streams in the Ari-

154 Endangered Species UPDATE Vol. 18 No. 4 2001 man population den- sity. Studies (e.g., Mladenoff et al. 1995; Ratti et al. 1999) have shown that lands with a hu- man population den- sity greater than 12 to 13 persons per square kilometer will not be suitable wolf habitat. The I' I I Figure 2. Vegetation cover Figure 3. Surface water I map displayed in Figure 6 indicates zona portion of the Grand Canyon teau enjoys a very high density of that most of the Arizona section of Ecoregion. Although the digital data mule deer (eight to 13 animals per the Grand Canyon Ecoregion has available is very incomplete, this fig- km2),while the remainder of the area population densities less than 13 per- ure shows that there are more than has an adequate density of three to sons per km2. Except for the Flag- enough sources of surface water on eight deer per km2. The Coconino staff-Sedona urban zone, the entire the Kaibab Plateau, in the Flagstaff Plateau around Flagstaff also supports north-south corridor from the Kaibab area, and along the Mogollon Rim. quite dense populations (three to eight Plateau to the eastern, slightly urban- This conclusion is supported by the per km2). Furthermore, similar den- ized, part of the Mogollon Rim has a observed presence of relatively high sities probably exist on parts of the human population density of less than numbers of other large predators (e.g., Hualapai and Navajo reservations in four people per km2. Not surprisingly, mountain lions (Felis concolor) and the GCE, but no data is readily avail- this same corridor has high prey spe- prey species in these locales. able to confirm this supposition. cies densities and seems capable of Clearly, one of the most impor- Even if the current mule deer popu- supporting wolves. tant determinants of suitable wolf lation density is one-half of what Wolves are usually not threatened habitat is the abundance of ungulate these figures indicate, as some Ari- by roads, except when they are struck prey species. The primary prey spe- zona Game and Fish Department per- by motor vehicles (Mech 1977). cies for wolves in the Grand Canyon sonnel suggest (e.g., Goodwin, per- Nonetheless, roads can provide ac- Ecoregion are mule deer (Odocoileus sonal communication),there are still cess to generally undisturbed areas hemionus), followed in order of im- more than sufficient deer densities to where humans may harass or kill portance by elk (Cervus elaphus), support gray wolves. Figure 5 shows wolves. Studies of road density and pronghorn (Antilocarpa americana), that elk densities, while somewhat wolf distribution relationships by and bighorn sheep (Ovis canadensis). lower on average than mule deer, are Thiel (1985) and Mech et al. (1988) Information about abundance (den- quite high (i.e. two to three animals suggest a road density threshold value sity) and distribution of these wild- per krn2) around Flagstaff and south- of between 0.6 and 0.8 kilometers of life species in the Arizona part of the east along the Mogollon Rim. Of road per square kilometer of area. ecoregion was obtained from the Ari- course, elk also average three times Higher road density values generally zona Game and Fish Department. the biomass of deer. Figures 4 and 5 result in unsuccessful breeding at- Figures 4 and 5 display the approxi- combined map an adequate ungulate tempts. Mladenoff et al. (1995), us- mate density of mule deer and elk prey base extending north-south from ing radio collar data on recolonizing populations in Arizona GCE. the Kaibab Plateau, through the Flag- wolves in northern Wisconsin, dis- Other reintroduction studies (e.g., staff area, and southeast along the covered that road density and fractal FWS 1996) indicate that a density of Mogollon Rim. dimension-reflecting the degree of approximately two to six deer per km2 habitat fragmentation (often the re- would be required to support a Mexi- Human dimensions sult of road buildingewere the most can wolf population and, presumably, An important determinant of habitat important predictors of favorable similar numbers would be adequate suitability for gray wolves and other wolf habitat. Figure 7 shows that for wolves in the GCE. Figure 4 dem- large carnivores such as grizzly bears most of the north-south corridor, ex- onstrates that much of the Kaibab Pla- (Merrill et al. 1999) seems to be hu- tending from the Kaibab Plateau

Vol. 18 No. 4 2001 Endangered Species UPDATE 155 through to the Mogollon Rim south- The landscape-scale east of Flagstaff, has road densities habitat mapping, included in higher than 0.68 krn per km2,but gen- this progress report, is adrnit- erally lower than 1.4 km per km2. tedly somewhat incomplete Road density in many parts of the at this stage in the research. GCE is somewhat higher than recom- Nonetheless, the mapped mended in other studies, but most of variables of both biophysical the numerous roads in the ecoregion and human dimensions point are tertiary or unimproved roads strongly towards the prob- that could be eliminated on public ability that at least two ar- lands with a vigorous road-closing eas-the Kaibab Plateau and program. Furthermore, the low hu- much of the Mogollon Rim man density numbers (Figure 6) south and east of Flag- Figure 4. Mule deer density might indicate that these areas are staff-are capable of sup- favorable wolf recovery habitat de- porting viable wolf popula- spite the existence of relatively high tions and suitable for rein- unimproved road densities. troduction of gray wolves Favorable land status, defined here as lands in public ownership and, Projected wolf densities especially, designated protected ar- Assuming that wolf reintro- eas, can help make a landscape suit- duction is feasible, it is rea- able for gray wolf reintroduction sonable to ask how many (Southern Rockies Ecosystem wolves might the Arizona Project, 1998). Identifying, describ- portion of the Grand Canyon ing, and mapping proposed and des- Ecoregion support. Utilizing ignated wilderness areas and other ar- the existing deer and elk den- eas designed to protect ecological sity distribution maps (Fig- Figure 5. Elk density processes or wildlife, such as the ures 4 and 5), and following Grand Staircase/Escalante National Fuller (1989), very prelimi- Monument and the Grand Canyon nary calculations of predicted Game Preserve in the Kaibab Forest wolf density were done us- (Miller 1996)is especially important. ing these equations: Figure 8 maps distribution of public lands, both state and federal, exclu- sive of Indian reservations. This re- veals that a wide band of federal pub- and lic lands (including large tracts of pro- tected areas) runs north-south from the Kaibab Plateau through the Flag- staff area and southeast along the where W is predicted wolf Mogollon Plateau (again, corre- density (per 10OCl km2), D is Figure 6. Human density sponding with the distribution of estimated mule deer density important biophysical factors). (per km2),and 3E is estimated State lands, even though currently elk density (per km2)times a relative suggest that to reintroduce at least 100 interspersed in a "checkerboard" biomass value (elk biomass is 3 x 1 gray wolves into the Arizona portion fashion (see Figure 8) with private deer). Both low andhighungulateden- of the Grand Canyon Ecoregion and federal lands, could be consoli- sity estimates were utilized in calculat- would be feasible. dated through land trades and pur- ing a range of predicted wolf numbers chases to create wildlife corridors shown in Table 1. Potential stock for wolf between federal public lands such Although this facet of the study reintroduction as the Coconino and Kaibab Na- is far from finished, historical records When the time comes to make a de- tional Forests. and initial carrying capacity research cision about reintroducing gray

156 Endangered Speclet? UPDATE Vol. 18 No. 4 2001 tundra) and experienced in tial reintroduction sites in the Arizona hunting the types of ungu- section of the Grand Canyon late species that are most Ecoregion. Initial results show that abundant in the GCE. there are at least two localities in Alternatively, captive or northern Arizona available for rein- wild-bred Mexican wolves troduction of around 100 wolves. (C. 1. baileyii) could be uti- Source stock for wolf recovery in the lized for reintroduction. GCE could come from existing large Given the difficulties expe- C. I. nubilus populations in the Great rienced with-captive bred Lakes andlor a recovered C. I. baileyii stock in the current Mexican population in the Southwest. Clearly, wolf recovery effort, how- the future extension of wolf recovery Figure 7. Road denisty ever, it seems best to wait for into northern Arizona and other parts the availability of surplus of the GCE will have to be done un- wild-raised stock GCE (par- der the legal mandate of the ESA sons, personal communica- and will most likely be sponsored tion). Also, when the Mexi- by a federal agency such as the Fish can wolf population reaches and Wildlife Service and/or Na- a viable size in the wild, dis- tional Park Service. persers from eastern Arizona Further investigation and study and western New Mexico will continue to refine the habitat ca- will likely attempt to colo- pability and suitability analyses, as nize the southeastern part of well as to help determine the most ap- the GCE. Thus, this recov- propriate subspecies for wolf reintro- ery opportunity in the GCE duction in the ecoregion. In the end, could help extend the geo- however, the most important consid- Figure 8. Land status graphic range and eration is how to best assist nature in metapopulation of the cur- restoring gray wolves to the Grand wolves to the Grand Canyon rently recovering, but still endan- Canyon Ecoregion and thereby help Ecoregion, we should " ...consider gered, Mexican wolf. in the national effort to conserve behavioral or demographic factors to this magnificent and ecologically be more important than maintenance Conclusions essential carnivore. of the genetic purity of putative wolf The first phase of this landscape-scale subspecies.. . " (Wayne et al. 1992). analysis involved utilizing six factors Acknowledgements If Nowak's (1995) recent revision of of the biophysical and human dimen- This project was primarily funded by wolf taxonomy is accepted, it seems sions to identify and describe poten- the Grand Canyon Wildlands Coun- biologically appropriate that stock for reintroduction could be taken from anywhere in the historic range of C. Table 1. Minimum and maximum number of wolves predicted to oc- I. nubilus. While finding areas of cupy Arizona part of Grand Canyon Ecoregion based on combined surplus wolf populations with habi- mule deer and elk densities and areas. tat exactly comparable to the GCE will be difficult, regions such as the Location Area (square kms) Wolves (pop. range) Great Lakes, currently supporting C.1. nubilus populations, do exhibit analo- gous forested ecosystems (albeit dif- North Kaibab 1472 35 to 62 ferent forest types) and have similar ungulate prey species (i.e., deer and South Colorado 3849 80 to 125 elk). Wild wolves translocated from Plateau the Great Lakes region, for ex- ample, would at least be habituated Total Arizona GCE 5321 115 to 187 to forest habitats (as opposed to

Vol. 18 No. 4 2001 Endangered Species UPDATE 157 cil, Defenders of Wildlife, and bana, Illinois. Final draft: feasibility study on the reintro- Prescott College. Although I am Hoffmeister, D.F. 1986. Mammals of Ari- duction of gray wolves to the Olympic Pen- solely responsible for any errors or zona. The University of Arizona Press: insula. Unpublished report prepared for the Tucson, AZ. United States Fish and Wildlife Service, shortcomings, I could not have made Lopez, B.H. 1978. Of wolves and men. Lacey, Washington. the progress I have without the help Charles Scribner's Sons, New York, New Russo, J.P. 1964. The Kaibab North deer of these people: Kelly Burke, Larry York. herd - Its history, problems and manage- Stevens, and Kim Crumbo of the Mech, L.D. 1970. The wolf: the ecology and ment: A research and management study, behavior of an endangered species. Uni- Wildlife Bulletin No. 7. Arizona Game and Grand Canyon Wildlands Council; versity of Minnesota Press, Minneapolis, Fish Department: Phoenix, Arizona. Melissa Grigione and Dan Smith, Minnesota. Sneed, P. and K. Crumbo. 1998. The rein- Defenders of Wildlife; Mike Mech, L.D. 1977. Productivity, mortality, troduction of gray wolves to the Grand Siedman, Phoenix Zoo; Mike Senn, and population trends of wolves in north- Canyon Ecoregion: A proposed feasibility Rick Miller, John Goodwin, Ray Lee, eastern Minnesota. Journal of Marnmology study. Unpublished research proposal. 58559-574. Grand Canyon Wildlands, Flagstaff, Ari- Matt Alderson, and Amber Munoz, Mech, L.D., S.H. Fritts, G.L. Raddel, and W.J. zona. Arizona Game and Fish Department; Paul. 1988. Wolf distribution and road Soule, M. and R. Noss. 1998. Rewilding and Elaine Leslie, Grand Canyon Na- density in Minnesota. Wildlife Society biodiversity: Complementary goals for con- tional Park; Rick Moore and Aimee Bulletin 16%-87. tinental conservation. Wild Earth 8(3): 18- Merrill, T., D.J. Mattson, R.G. Wright, and 28. Porcaro, Grand Canyon Trust; Dave H.B. Quigley. 1999. Defining landscapes Southern Rockies Ecosystem Project. 1998. Hanna and Craig Martinsen, Prescott suitable for restoration of grizzly bears Wolf recovery in the Greater Southern College; and, Jim Riedman, Arizona Ursus arctos in Idaho. Biological Conser- Rockies: an initial suitability analysis. Land Resources Information System. vation 87:23 1-248. Unpublished in-house report, Nederland, Miller, R. 1996. History of the intent and Colorado. management of the Grand Canyon Game Terbourgh, J., J.A. Estes, P. Paquet, K. Ralls, Literature eited Preserve. Unpublished report prepared for D. Boyd, B. Miller and R. Noss. 1999. The Brown, D.E., ed. 1984. The wolf in the south- Arizona Game and Fish Department, Flag- role of top carnivores in regulating terres- west: the making of an endangered species. staff, Arizona. trial ecosystems. Pp 39-64 in Soule, M. The University of Arizona Press, Tucson, Mladenoff, D.J., T.A. Sickley, R.G. Haight, and J. Terbourgh, ed. Continental conser- Arizona. and A.P. Wydeven. 1995. A regional land- vation: design and management principles Clark, T.W., A.P. Curlee, and R.P. Reading. scape analysis and prediction of favorable for long-term, regional conservation net- 1996. Crafting effective solutions to the gray wolf habitat in the northern Great works. Island Press: Washington, D.C. large carnivore conservation problem. Lakes region. Conservation Biology 9:279- Thiel, R.P. 1985. Relationship between road Conservation Biology 10:940-948. 294. densities and wolf habitat suitability in Fuller, T.K. 1989. Population dynamics of Nowak, R.M. 1995. Another look at wolf Wisconsin. American Midland Naturalist wolves in north-central Minnesota. Wild- taxonomy. Pp 375-398 in L.N. Carbyn, 113:404-407. life Monographs 105:1-41. S.H. Fritts, and D.R. Seip, eds. Ecology United States Fish and Wildlife Service. 1996. Fuller, T.K., G.L. Raddle, M.S. Lenarz, and and conservation of wolves in a changing Final environmental impact statement- G.B. Joselyn. 1992. A history and current world. Occasional Publication No. 35, Reintroduction of the Mexican wolf within estimate of wolf distribution and numbers Canadian Circumpolar Institute, University its historic range in the southwestern United in Minnesota. Wildlife Society Bulletin of Alberta, Edmonton, Alberta. States. U.S. Fish and Wildlife Service, 20:42-55. Paquet, P.C. and A. Hackman. 1995. Large Albuquerque, New Mexico. Gipson, P.S., W.B. Ballard, and R.M. Nowak. carnivore conservation in the Rocky Moun- Wayne, R.K., N. Lehman, M.W. Allard, and 1998. Famous North American wolves and tains. World Wildlife Fund Canada, R.L. Honeycutt. 1992. Mitochondria1 the credibility of early wildlife literature. Toronto, Ontario. DNA variability of the gray wolf: genetic Wildlife Society Bulletin 26:808-8 16. Quinby, P., S. Trombulak, T. Lee, J. 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158 Endangered Species UPDATE Vol. 18 No. 4 2001 Canis Soupus: Eastern Wolf Genetics and Its Implications for Wolf Recovery in the Northeast United States

Nina Fascione Defenders of Wildlife, 1101 14th St. NW, Suite 1400, Washington DC 20005; [email protected] Lisa G. L, Osborn Defenders of Wildlife, P.O. Box 756, Shelburne, VT 05482; [email protected] Stephen R. Kendrot USDNAPHIS/Wildlife Services, P.O. Box 130, Hull Street Rd., Mosely, VA 23120; [email protected] Paul C, Paquet Faculty of Environmental Design, University of Calgary, Calgary, AB, Canada; ppaquet0sk.syrnpatico.ca

Abstract Efforts to restore wolves to the northeastern United States have been confounded by a new taxonomic proposal: that the wolf historically inhabiting this region was not, as previously thought, a subspecies of gray wolfcommonly called the eastern timber wolf(Canis lupus lycaon), but rather a separate species closely related to the red wolf(Canis rufus) of the southeast United States. This hypothesis raises numerous biological, legal, policy, and management questions about potential wolf restoration. While restoring wolves could complete a broken food chain by providing a natural predator for moose in the northern forest ecosystem, the process of wolf restoration in the Northeast is in its infancy. Further studies must address biological, sociologi- cal, and economic impact questions, as well as answer the basic question of what wolf originally inhabited the northeastern forests?

Introduction Species Act (ESA), the U.S. Fish and this region. If enacted, this designa- Efforts to restore wolves to the north- Wildlife Service (FWS) listed east- tion would separate the Northeast eastern United States have been con- ern timber wolves as endangered, administratively under the ESA from founded by a new taxonomic pro- except in Minnesota where a remnant wolf populations in the Great Lakes posal: that the wolf historically in- population was listed as threatened. states and require the Service to de- habiting this region was not, as pre- In 1978, the Service developed a re- velop a new recovery plan for New viously thought, a subspecies of gray covery plan for the eastern timber England and upstate New York. Re- wolf commonly called the eastern wolf. At that time, scientists believed cent studies indicate there are ad- timber wolf (Canis lupus lycaon),but that the eastem timber wolf had his- equate habitat and prey to sustain a rather a separate species closely re- torically ranged throughout the healthy wolf population in this region lated to the red wolf (Canis rubs) of Northeast and west to the Great Lakes (Harrison and Chapin 1997; the southeast United States (Wilson region. The recovery plan identified Mladenoff and Sickley 1998), and et al. 2000). This hypothesis raises several areas in the Northeast as po- several surveys indicate strong pub- numerous legal, policy, and manage- tential wolf habitat, including north- lic support for wolf restoration (Re- ment questions about potential wolf west Maine and the Adirondack sponsive Management 1996; Downs restoration. In addition, wildlife Mountains of New York. These ar- and Smith 1998). However, the un- managers now have a basic biologi- eas remained in the recovery plan resolved issue of taxonomy, while cal question to consider when debat- when it was revised in 1992 but the certainly not the only impediment to ing the merits of wolf reintroduction Service did not actively pursue north- northeast wolf restoration, is compli- to New England and upstate New east wolf restoration. cating the prospect. York: what wolf should be restored? A recent proposal by the Service Wolves were extirpated from the to declare a Distinct Population Seg- Previous taxonomic Northeast by the end of the nineteenth ment for wolves in the Northeast classification century (Fowler 1974). In 1974, a (Federal Register 2000) has triggered In this article, we use the term east- year after passage of the Endangered renewed interest in wolf recovery in em wolf to refer to the wolf that, by

Vol. 18 No. 4 2001 Endangered Species UPDATE 159 Nowak's description (1995), cur- and fragmentation favored their east- monly found west of Algonquin Park, rently resides in southeastern Canada ward expansion (Clarke 1970; Wayne whereas the park itself maintains the and formerly inhabited the northeast- and Lehman 1992; Roy et al. 1994). most wolf-like form of eastern wolf ern United States. We also use the In the 1920s, coyotes moved east (Wilson et al. 2000). term western gray wolf to refer to across southern Ontario and east- The result of these genetic analy- what Nowak (1995) describes as Ca- ern Quebec and by the 1930s had ses is the discovery of a canid gradi- nis lupus nubilus, the larger wolf that reached New York and New En- ent in the northeastern United States formerly inhabited much of the west- gland (Parker 1995). The south- and southeastern Canada, containing ern United States and much of eastern United States similarly un- a mix of eastern wolf, western gray Canada. Goldman (1937) classified derwent landscape changes that wolf and coyote genes. This phe- the eastern wolf as Canis lupus enabled coyotes to expand to this nomenon is often lightheartedly re- lycaon, a subspecies of gray wolf, and region as gray and red wolves were ferred to as "canid soup." According for years its historic range was extirpated in the last century (Jenks to Wilson et al. (2000), in the north- thought to be the northeastern United and Wayne 1992; Parker 1995). eastern United States, coyote genes States as far west as the Great Lake As coyotes expanded eastward, dominate this mix. 1n southeastern states and north into southern Ontario they encountered dwindling popula- Ontario, eastern wolf genes are more and Quebec (Goldman 1944; FWS tions of both eastern and red wolves. common, and in northeastern and 1992; Nowak 1995). The red wolf is There has been much discussion in northwestern Ontario an eastern classified as Canis rufus, a distinct the literature about the propensity for wolflwestern gray wolf animal may species (Goldman 1937), and its and degree of interbreeding between be predominant. The extent to which historic range has long been con- red wolves and coyotes, both earlier eastern wolves interbreed with west- sidered to be the southeastern this century and since their reintro- ern gray wolves is currently un- United States as far west as Texas duction in North Carolina (Wayne known. The regions in far northern and as far north as Pennsylvania and Jenks 199 1; Nowak 1992; Kelly Ontario are predicted to be predomi- (Nowak 1995). By these boundary et al. 1999 ). At present, the FWS is nantly western gray wolf, but at definitions, the ranges of the red struggling to keep reintroduced red present the boundary between the wolf and eastern wolf would have wolves from hybridizing with coy- eastern wolf and western gray wolf originally overlapped in the mid- otes. If the current rate of interbreed- is not well established (P. Wilson, Atlantic states (Nowak 1995), in- ing is not halted, red wolf genes will personal communication). cluding Pennsylvania, West Vir- be completely diluted within a few ginia, and Virginia (see Figure 1). generations in a process known as Genetic hypothesis Over the years, scientists noted genetic swamping (Kelly et al. 1999). The tendency for eastern and red similarities between the two canids Genetic testing of the relatively wolves to hybridize with coyotes is based on morphology and skull large coyotes from the Adirondack not observed in western gray wolves measurements (Lawrence and Park and central New York similarly (Edwins et al. 2000; Wilson et al. Bossert 1975; Alexander 1983 ). indicates a history of interbreeding 2000). Additionally, there are mor- The two animals are apparently so with wolves. The degree of wolf ge- phological characteristics shared by similar that one report in the litera- netic material varies across these red and eastern wolves but not west- ture 25 years ago refers to a red samples, with some being more ern gray wolves, such as the smaller wolf in Algonquin Park, Ontario (P. " wolf-like" than others (Chambers size of wolves in the East (Goldman Wilson, personal communication), 2000). Genetic testing on northern 1944). These commonalities led re- even though scientists did not think New England canids shows inter- searchers to examine more closely they ranged that far north. breeding as well, though more sam- the relationship between red wolves pling is needed. In the Frontenac and eastern wolves. They hypoth- Recipe for canid soup Axis region of Ontario, southeast of esized that red and eastern wolves Scientists noted another trait com- Algonquin Park, a similar, though were more closely related to each mon to both red wolves and eastern slightly larger canid is commonly other than either was to western gray wolves-the tendency to hybridize called the Tweed wolf, and is prob- wolves (Wilson et al. 2000). In the with coyotes, Canis latrans (Wilson ably a hybrid containing more wolf past decade, researchers from the et al. 2000). Coyotes, historically ab- genes than coyote genes (Edwins et genetics labs at Trent and McMasterts sent from the east, reached Ontario al. 2000; P. Wilson personal commu- universities in Ontario have con- in the early 1900s as habitat alteration nication). A coyote-wolf mix is com- ducted genetic analyses of canids

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I60 Endangered Species UPDATE Voi. 18 No. 4 2001 Discussion If the issue of eastern canid taxonomy is the subject of debate for academi- cians, it is nothing less than con- founding to the public, legislators and even wildlife managers, who want to make informed decisions about wolf restoration in the Northeast. The con- cept that the eastern wolf and red wolf are the same species complicates an already complex topic, presenting nu- merous questions to all those inter- ested in examining the potential for wolf recovery in New England and New York. What canid originally oc- cupied the Northeast and what canid is there now? What role is the present canid filling and what role would a restored, larger canid fill? What wolf population would scientists use as source animals, and would they dis- place or interbreed with the Northeast's resident coyotes? Because of the tendency of the eastern wolf to hybridize with coy- otes, it is a fair assumption that if it were reintroduced in the Northeast it Figure 1. Historic and current ranges of the red and eastern wolf would be vulnerable to interbreeding with eastern coyotes. Therefore, de- spite efforts to reintroduce a separate from throughout the red and eastern land bridge and evolved into the species in the Northeast, biologists wolf range, using various criteria to gray wolf. The remaining canids might just be regenerating an animal compare relatedness. Their data sup- evolved wholly in North America. that is already present in the form of port the hypothesis that the red wolf Between 150,000 and 300,000 a large, hybrid coyote. It is impor- and eastern wolf have. a common years ago they diverged into the tant to note, however, that the core North American origin separate from coyote, which adapted to preying population of eastern wolves within that of the western gray wolf. In fact, on smaller mammals in the arid Algonquin Park is not readily hybrid- these researchers suggest that the red southwest, and the eastedred wolf, izing with coyotes, although the rea- wolf and eastern wolf are actually the which adapted to preying on white- son for this is unclear (Edwins et al. same species, and they propose tailed deer (Odocoileus 2000). Habitat saturation by estab- changing the scientific name of both virginianus) in eastern forests. lished wolf packs might be one ex- to Canis lycaon, with a common Gray wolves returned to the North planation. Protected wolves that are name of red wolf (Wilson et al. 2000). American continent approximately able to maintain long-term territories The recommended name is based on 300,000 years ago, adapting to are able to prevent the encroachment historical precedence in early wolf preying on large ungulates through- of coyotes into core wolf areas (J. taxonomy (Brewster and Fritts 1995). out the western United States and Theberge, personal communication). According to Wilson et al. Canada. According to this hypoth- Canadian wolf researcher John (2000), North America was inhab- esis, coyotes are more closely re- Theberge has documented short-term ited by a common canid ancestor lated to the easternlred wolf than to invasion by coyotes following the one to two million years ago. At the western gray wolf, hence the breakup of a resident wolf pack some point, some of these animals propensity for interbreeding (Wil- within Algonquin's core wolf zone (J. traveled to Eurasia over the Bering son et al. 2000). Theberge, personal communication).

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Vol. 18 No. 4 2001 Endangered Species UPDATE 161 Understanding this relationship could environment, indicating that perhaps that the Laurentides animal is the prove useful in the Northeast to de- both canids-the larger gray wolf and most suitable for Northeast reintro- termine what conditions are neces- the smaller eastern wolf-might have duction, but cannot definitively sary to discourage interbreeding. inhabited the northeast at various prove its earlier existence there, From an ecological perspective, points (P. Wilson, personal commu- would the public accept a species wildlife managers will have to deter- nication). Given the radical changes whose original range may have mine what the most appropriate spe- that have occurred in the Northeast stopped short of the proposed rein- cies of wolf is for the modem day ecosystem since colonial times, and troduction area? These types of moose-dominated North Woods. the lack of remaining physical evi- debates will most certainly slow the Should a species' historical presence dence of the presence of wolves, it is process of wolf restoration. be the determining factor, or should difficult to determine which species the most suitable candidate be se- may have been present. Conclusion lected to match the ecological condi- Similarly, some scientists specu- While the current genetics research tions existing today or likely to exist late that the eastern wolf did not his- is intriguing, and increasingly gain- in the future? If so, the wolves in the torically occur north of the Canadian ing the support of the scientific com- Laurentides area of Quebec might border (P. Wilson, personal commu- munity (Kelly, personal communica- provide a better source population for nication). Southeastern Canada was tion), further research and discussion the northeastern United States. The home to moose, woodland caribou is essential before conclusions can be close proximity of the two regions (Rangifer tarandus caribou) and elk drawn. In the meantime, other bio- might allow some genetic exchange. (Cewus elaphus) and would have logical factors are ripe for wolf res- Preliminary testing indicates that ge- more likely contained the larger gray toration. The northern forests have netically, the Laurentides animals wolf. Overhunting and habitat alter- made a remarkable recovery in the may be a mix of gray and eastern wolf ation likely contributed to the decline last fifty years and now comprise (P. Wilson, personal communication), of some of these species, including more than 26 million acres-ample a finding which is further supported the wolf (Nelson 1997). As intensive habitat to support a population of by their large size of up to 100 logging encroached into southeastern wolves (Harrison and Chapin 1998; pounds. The Laurentides wolves Canada, white-tailed deer popula- Mladenoff and Sickley 1998). With prey primarily on moose (Alces tions expanded northward from the the expansion of moose and beaver alces), a plentiful prey item in Maine. United States, thriving in the second (Castor canadensis) and the occur- Most importantly perhaps, these ani- growth forests that resulted. Eastern rence of white-tailed deer throughout mals may be less likely to hybridize wolf populations may have likewise the north woods, there is ample prey. with eastern coyotes than the moved north in an expansion similar Most scientists agree that there is an Algonquin-type wolf, though this to their primary prey species. Today, ecological role for a larger canid in crucial relationship remains to be the range of the eastern wolf may the northeast's North Woods tested (P. Wilson, personal commu- extend as far west as Saskatchewan (Harrison quoted in Fascione and nication). In the world of survival of and include northwestern Ontario and Kendrot, 2000). Even though coy- the fittest, it is unlikely that natural Minnesota. These northern areas otes occasionally form packs, they do selection would favor a moose-eat- may contain both eastern wolves and not normally prey on moose. Restor- ing canid that compromises its size western gray wolves, but more re- ing wolves could complete a broken by breeding with the much smaller search is needed before this can be food chain by providing a natural eastern coyote. determined (Wilson et al. 2000). predator for moose in the northern This theorizing, however, begs The sociological implications of forest ecosystem. Since the 1995 the question: did this larger gray wolf the new taxonomic proposal are as wolf reintroduction in the northern ever exist in the Northeast? For now, challenging as the biological. Wolf Rockies, the Greater Yellowstone scientists can only speculate, but it is restoration is always politically divi- ecosystem has undergone significant generally accepted that the northeast- sive. Groups and individuals oppos- ecological changes as a result of the ern United States was primarily a ing wolf restoration have already restoration of this top predator. Recent moose-caribou ecosystem before Eu- latched onto the current debate, de- research in Yellowstone National Park ropean settlement (D. Harrison, per- claring that the role of the eastern suggests that the effect wolves have on sonal communication). It is question- wolf is filled by the large, resident their prey can benefit vegetative struc- able whether the deer-adapted east- eastern coyote, which many call the ture and overall species biodiversity ern wolf would have thrived in this "brush wolf." If scientists determine (Ripple and Larsen, in press).

162 Endangered Species UPDATE Vol. 18 No. 4 2001 The process of wolf restoration published manuscript. the northeastern United States: a spatial in the Northeast is in its infancy, how- Clarke, C.H.D. 1970. Temporary Study Com- prediction of favorable habitat and poten- mission on the Future of the Adirondacks. tial population levels. Journal of Wildlife ever, and further studies must address Technical Report a. Management 62: 1-10. biological, sociological and eco- Downs, G. and L. Smith. 1998. Summary Nelson, R. 1997. Heart and blood: living with nomic impact questions (Fascione et Report of the Maine Wolf Attitudes Sur- deer in America. Random House, Inc. New al. 2000). While the proposed taxo- vey. Center for Research and Evaluation. York, NY. nomic revision highlights the need to University of Maine, Orono. Nowak, R.M. 1992. The red wolf is not a hy- Edwins, P., M. de Almeida, P. Miller and 0. brid. Conservation Biology 6(4):593-595. identify the connections and poten- Byers, eds. 2000. Population and Habitat Nowak, R.M. 1995. Another look at wolf tax- tial interactions between eastern Viability Assessment Workshop for the onomy. Pp 375-397 in L.N. Carbyn, S.H. wolves, red wolves, gray wolves and Wolves of Algonquin Park: Final Report. Fritts and D.R. Seip, eds. Ecology and con- eastern coyotes in the Northeast, a IUCNISSC Conservation Breeding Spe- servation of wolves in a changing world: cialist Group: Apple Valley, MN. proceedings of the second North Ameri- clearer picture of these complex re- Fascione, N., 0. Byers, U.S. Seal, eds. 2000. can symposium of wolves. Canadian Cir- lationships is not likely to simplify Wolves in the Northeast? A Workshop to cumpolar Institute, University of Alberta, the management, policy, legal, and Exchange Views. Final Report. IUCNISSC Edmonton, Canada. moral issues that follow. Although Conservation Breeding Specialist Group: Parker, G. 1995. Eastern Coyote: the story the questions raised by this taxo- Apple Valley, MN. of its success. Nimbus Publishing. Halifax, Fascione, N. and S. Kendrot. 2000. The Nova Scotia. nomic discussion are unique to the Northeastern wolf question. Defenders Responsive Management. 1996. Public Opin- Northeast, controversy itself is noth- Winter 2000/2001:27-33. ion on and Attitudes toward the Reintro- ing new to wolf restoration. Ulti- Federal Register. July 13, 2000. 65(135). duction of the Eastern Timber Wolf to mately, genetics are just one more Fowler, B. 1974. Adirondack Album. Out- Adirondack Park. Presented to Defenders door Association, Schenectady, New York. of Wildlife, Washington, DC. piece of information that wildlife Goldman, E.A.1937. The wolves of North Ripple, W.J., and E. J. Larsen. In Press. His- managers, advocates, and the public America. J. Mammalogy 18:37-45. toric aspen recruitment, elk, and wolves will have to distill and blend with the Goldman, E.A. 1944. Classification of in N. Yellowstone National Park, USA. ecological and sociological realities wolves. Pp 389-636 in S.P. Young and Biological Conservation. of not only today's environment, but E.A. Goldman. The wolves of North Roy, M.S., E. Geffen, D. Smith, E.A. America. American Wildlife Institute, Ostrander, and R. K. Wayne. 1994. Pat- tomorrow's as well. Washington, D.C. terns of differentiation and hybridization Harrison, D.J., and T.G. Chapin. 1997. An in North American wolflike canids, re- Acknowledgments assessment of potential habitat for eastern vealed by analysis of microsatelitte loci. The authors are very grateful to Paul timber wolves in the northeastern United Molecular Biology and Evolution States and connectivity with occupied 11553-570. Wilson, not only for reviewing this habitat in southeastern Canada. A sum- U.S. Fish and Wildlife Service. 1992. Recov- manuscript, but for always being mary report and position paper prepared ery Plan for the eastern timber wolf. Twin willing to come out of the lab to ex- for the Wildlife Conservation Society. Cities, Minnesota. plain this research. We would also Working Paper No. 7, March 4, 1997. Wayne, R.K. and S. Jenks. 1991. Mitochon- like to thank Paul Nickerson, Michael Jenks, S.M. and R.K. Wayne. 1992. Prob- drial DNA analysis implying extensive lems and policy for species threatened hybridization of the endangerd red wolf Amaral, Bob Ferris and Martin Smith by hybridization: the red wolf as a case Canis rufus. Nature 351565568, for their review and feedback. study. Pp 237-25 1 in McCullough, D.R., Wayne, R.K. and Lehman.1992. Mitochon- and R.H.Barrett, eds. Wildlife 2001: drial DNA analysis of the eastern coyote: Literature cited Populations. New York: Elsevier Ap- origins and hybridization. Pp 9-22 in A.H. Alexander, H.E. 1983. The red wolf saga. plied Science. Boer, ed. Ecology and management of the Arkansas Game & Fish 14(2): 2-6. Kelly, B.T., P.S. Miller and U.S. Seal, eds. eastern coyote. Wildlife Research Unit, Brewster, W.G. and S.H. Fritts. 1995. Tax- 1999. Population and Habitat Viability 'University of New Brunswick, onomy and genetics of the gray wolf in Assessment Workshop for the Red Wolf Fredericton. North America: a review. Pp 353-373 in (Canis rufus). Apple Valley, MN: Con- Wilson, P.J., Grewal, S., Lawford, I.D., L.N. Carbyn, S.H. Fritts and D.R. Seip, servation Breeding Specialist Group Heal, J.N.M., Granacki, A.G., Pennock, eds. Ecology and conservation of wolves (IUCNISSC). D. Theberge, J.B., Theberge, M.T., in a changing world: proceedings of the Lawrence, B. and W.H. Bossert. 1975. Rela- Voigt, D.R., Waddell, W., Chambers, second North American symposium of tionships of North American Canis shown R.E., Paquet, P.C., and Goulet, G., Cluff, wolves. Canadian Circumpolar Institute, by a multiple character analysis of selected D. and B.N.White. 2000. DNA profiles University of Alberta, Edmonton, Canada. populations. Pp 73-86 in M.W. Fox, ed. of the eastern Canadian wolf and the red Chambers, R.C. 2000. Taxonomic and di- The wild canids. Van Nostrand Reinhold, wolf provide evidence for a common etary characteristics of eastern coyotes New York. evolutionary history independent of the (Canis latrans var.) in New York state. Un- Mladenoff, D.J. and T.A. Sickley. 1998. As- gray wolf. Canadian Journal of Zoology sessing potential gray wolf restoration in 78:2156-2166.

Vol. 18 No. 4 2001 Endangered Species UPDATE 163 Captivity, Inbreeding, Cross-Lineage Matings, and Body Size in Mexican Wolves Richard Fredrickson Department of Biology, Arizona State University, Tempe, AZ 85287-1501; [email protected] Philip Hedrick Department of Biology, Arizona State University, Tempe, AZ 85287-1501; [email protected]

Abstract The Mexican gray wolf (Canis lupus baileyi) appears to be extinct in the wild and exists now only in captivity and in a single, small, reintroduced population. A recent study of captive animals found no evidence for inbreeding depression in juvenile viability or litter size. We examined the relationship between inbreeding and body weight in captive wolves. We found that captive wolves with little or no known inbreeding had lower body size than wild caught wolves. In addition, captive wolves with higher inbreeding had lower body size than captive wolves with little or no inbreeding. The captive population was descended from three founders until two other lineages, each descended from two founders, were recently added to the population. Consequently we examined the potential sfatistical power associated withfiture comparisons of body weights between inbred wolves and the offspring of cross-lineage matings. In this case it appears likely that in the next few years there will be an adequate sample size to statistically evaluate differ- ences in body size between these groups.

Introduction current population of wolves origi- which a substantial amount of data Minimizing inbreeding and the loss nated from three independent captive currently exists for captive Mexi- of genetic variation are major con- lineages with each having only two can wolves is adult body size mea- cerns of captive breeding programs or three founders captured during the sured by mass. Some body size for endangered species. Increases 1960s and 1970s from northern data is also available from histori- in inbreeding have been correlated Mexico and southern Arizona. Be- cal wild-caught wolves (see below). with decreases in juvenile survival cause of the few founders, each of the For these reasons and others, body in captivity in a number of differ- lineages has become inbred in cap- size has been proposed as a pheno- ent species (Ralls et al. 1988). In tivity. Following a genetic evalua- typic indicator with which to moni- addition, a number of recent stud- tion of the three lineages (Hedrick et tor the effects of merging the three ies in wild po,pulations have dem- al., 1997) in which no evidence of lineages. Because until recently onstrated that a reduction in genetic coyote, (Canis latrans), or dog, (Ca- there had been no breeding between variation resulting from inbreeding nis familiaris), ancestry was found, the three lineages, offspring from andlor genetic drift has been asso- the McBride, Ghost Ranch, and crosses between lineages will be ciated with the decay of phenotypic Aragon lineages were recently com- free of inbreeding. If inbreeding attributes related to fitness in wild bined to form a single population has negatively affected body size or populations (see references in with seven founders. other fitness related traits then in- Hedrick and Kalinowski 2000). The most common phenotypic dividuals free of inbreeding may Mexican wolves (Canis lupus trait used to evaluate the effects of show changes in these traits. Ex- baileyi), the southern-most subspe- fitness in captive populations of en- cept for a few offspring that have re- cies of gray wolf, currently exist pri- dangered species has been juvenile sulted from recent crosses between marily as a captive population num- survival; secondarily, reproductive the lineages, all Mexican wolves bering about 200 individuals. Other traits, such as litter size, have been alive today are inbred to some extent. than a small, recently reintroduced used. Kalinowski et al. (1999), For wolves, changes in body population in Arizona and New however, recently found no effect size resulting from inbreeding in Mexico, there have been no con- of inbreeding on juvenile survival captivity may result in reduced firmed sightings of wild Mexican and litter size in captive Mexican fitnesses of individuals reintro- wolves in more than 20 years. The wolves. The only other trait for duced into the wild. This may con-

164 Endangered Specles UPDATE Vol. 18 No. 4 2001 sequently result in a lower viabil- sons involving captive wolves, we and female wolves were considered ity of wild, reestablished popula- used 71 masses from 26 male and separately in all analyses. Com- tions because body size is impor- 24 female adult wolves from the parisons between groups were tant in successful prey capture. McBride captive lineage. Two of made using t and nonparametric Further, if body size has been af- these wolves were wild-caught, and Mann-Whitney U tests. fected by inbreeding, other fitness- one was a lineage founder. Only related traits may also be adversely masses from wolves 1.5 to 9.5 years Results and discussion affected. Besides inbreeding, the old that were taken from August Comparisons using existing data captive environment and its related through January were used. Of the For wild and captive wolves com- husbandry may also potentially re- three lineages, only the McBride bined, female body sizes ranged sult in changes in fitness-related lineage was managed from its in- from 20.2 to 37.4 kg, while male traits by exerting selective forces dif- ception as part of a captive breed- sizes ranged from 21.8 to 41.3 kg. ferent from the natural environment. ing program; the weight data used Historic wild male and female Here we look for evidence of here was taken during the course of wolves were significantly larger an effect of inbreeding on adult these activities. Because the other than wolves in the McBride lineage body size in captive Mexican lineages were not similarly man- with little or no inbreeding (fe- wolves and evaluate the utility of aged until more recently, similar males: t = 4.01, df =24 , P< 0.001; this trait for detecting potential fu- data for the Ghost Ranch and males: t = 4.24, df = 23, P< 0.001). ture changes in size in offspring Aragon lineages do not exist. Even with the limited data currently from cross-lineage matings. Spe- Because the level of inbreeding available, the mean size of inbred cifically we ask (1) whether historic in the captive wolves varied we female wolves (f > 0.125) was sig- wild-caught adult wolves were split the wolves into two groups nificantly smaller than lhose of fe- larger than captive adult wolves based on the level of inbreeding in males with little or no inbreeding with little or no inbreeding and (2) individuals. Wolves with no in- (Mann-Whitney U=112.00, P< whether wolves in the McBride breeding in captivity (inbreeding 0.01), however, there was not a sig- captive lineage with little or no in- coefficient f = 0) and wolves com- nificant difference between males breeding were larger than adults prising the first generation of inbred (t=1.18, df=24, P=0.129) (Table 1). later in the lineage with greater individuals (f = 0.125) were com- ' The mean difference in body size amounts of inbreeding. In addition, bined to form a group with little or for females in the two groups is we examine the statistical power as- no inbreeding, and wolves with only 3.0 kg (about an 11% size re- sociated with future comparisons greater levels of inbreeding (f > duction), a value that becomes sig- when larger samples of inbred 0.125) were combined to form the nificant primarily because of the and cross-lineage wolves become second group (Table 1). The maxi- low variance in body mass in inbred available to assess the likelihood mum inbreeding coefficient in our females (2.4). The comparison be- of detecting differences associ- sample was 0.25. Because body tween males had only a 30% chance ated with inbreeding and out- size varies between the sexes, male of detecting a difference at the a= breeding, should they occur.

Materials and methods Table 1. Sample sizes and mean body sizes (kg) for historic, wild For comparisons involving historic, Mexican wolves and captlve wolves by level of inbreeding. wild wolves we used the masses of 28 wild Mexican wolves killed in Chihuahua and Durango, Mexico during the 1970s as part of control efforts (McBride, 1980, as reported in Brown, 1983). Full masses were reported for only eight of the 20 wolves; the remaining wolves had their viscera removed before weighing. Sizes of gutted wolves were adjusted to approximate their full weight (Table 1). For compari-

Vol. 18 No. 4 2001 Endangered Species UPDATE 165 0.05 level if a difference truly ex- gest that differences between males provided by the US Fish and Wild- ists, given the means, variances and with little inbreeding and those with life (RJF) and the Ullman Distin- sample sizes of the two groups. greater inbreeding will probably not guished Professorship (PWH). S. Although inbreeding did not be detectable with the sample sizes Kalinowski provided helpful com- appear to have an effect on viabil- attainable in the next few years ments and C. Buchanan, P. Miller, ity and litter size in Mexican wolves without a lower mean mass or vari- and P. Siminski provided the data (Kalinowski et al., 1999), our re- ance in the larger sample of inbred used in this work. sults provide some evidence of wolves. Differences, however, be- changes in body size that may be tween inbred wolves and the off- Literature cited manifestations of inbreeding de- spring from cross-lineage matings Brown, D.E. 1983. The wolf in the south- pression. Conclusions from these will likely be detectable, should west, the making of an endangered spe- cies. Univ. of Arizona Press, Tucson. small samples, however, may or they occur, with the samples sizes Hedrick, P.W. and S.T. Kalinowksi. 2000. may not be supported when larger likely available in the next few Inbreeding depression in conservation samples from inbred wolves in the years. In other words, within the biology. Ann. Rev. Ecol. Syst. 31:139- McBride lineage become available. next few years, assuming several 162. Such samp1e.s would he helpful in litters of cross-lineage wolves are Hedrick, P.W., P.S. Miller, E. Geffen, and R. Wayne. 1997. Genetic evaluation of determining more definitively produced per year, enough data the three captive Mexican wolf lineages. whether there is any evidence of a should soon be available to evalu- Zoo Biology 16: 47-69. decline in size among inbred ate the effect of merging the lin- Kalinowski, S .T., P. W. Hedrick, and P.S. wolves. Masses from additional eages on body size. It will be in- Miller. 1999. No evidence of inbreeding depression in Mexican and red wolves. wolves with little or no inbreeding teresting to determine if crossing to Conservation Biology 13: 137 1- 1377. in the McBride lineage will not be other lineages will genetically re- McBride, R.T. 1980. The Mexican wolf available because most of these in- store fitness-related traits in Mexi- (Canis lupus baileyi). US Fish and dividuals are dead and no such can wolves, as found in recent stud- Wildlife Service. Endangered Species matings are now possible. ies of other species. Report 8: 1-38. Ralls, K., J.D. Ballou, andA.R. Templeton. 1988. Estimates of lethal equivalents and Comparisons using future data Acknowledgements the cost of inbreeding in mammals. Con- Analyses of statistical power sug- Financial support for this work was servation Biology 2: 185-193.

166 Endangered Species UPDATE Vol. 18 No. 4 2001 First Swift fox, Vulpes velox, Reintroduction in the USA: Results of the First Two Years

Clio Smeeton Cochrane Ecological institute, PO Box 484, Cochrane, AB, Canada TOL OWO; (403) 932-5632; (fax) (403) 932-6303; cei Qcadvision.com Ken Weagle CEI Consulting Ltd., 7109 Bulyes Ave., Fort McMurray, AB, Canada T9H 5J6; kweagleQhome.com

Abstract The swift fox, Vulpes velox, is native only to the great plains of North America. The species was completely eradicated from its Canadian range by 1938, andfrom over 90% of its range in the USA by 1993. There have been two attempts at reintroducing the swift fox in to its historic range, one in Canada (1983 to 1997) and one on Blackfeet Tribal Land in the USA (1998 topresent). The reintroduction of the swift fox to the Blackfeet Tribal Lands in Montana began in August 1998. This is thefirst swift fox reintroduction to be undertaken in the USA. The project is afive- year partnership between the Blackfeet Fish and Wildlife Department, Cochrane Ecological Institute and Defenders of Wildlife. The reintroduction methodology used was based on that developed by the Cochrane Ecological Institute. A total of 76 swift fox have been released to date, and two years remain in the program. Survival of released animals is still being investi- gated; survival of radio collared swift fox released in 1999 was 75% over a one-yearperiod. Cubs have been produced in all years of the program. More than 24 cubs were observed in the field in 1999 and 2000.

Introduction of food per day, in the absence of fox had vanished throughout its Ca- Two species of fox, the water they absorb their needed liq- nadian range and from much of its (Vulpes macrotis) and the swift fox uid from their food and require 330 northern range in the U.S. reducing (Vulpes velox), are native to the great g of food per day (Flaherty and the possibility of subspecies verifi- plains of North America. Histori- Plaake 1986). Swift foxes are preyed cation. Definitive research, combin- cally, the swift fox range coincided upon by both terrestrial and avian ing morphological and molecular with that of the North American bi- predators and are extremely den de- systematic data (Stromberg and son, Bison bison. The species was pendent. Swift foxes are sedentary, Boyce 1986; Wayne1998) has re- once commonly found over a vast extremely social and largely monoga- sulted in the acceptance of distinct area, ranging from the eastern slopes mous. They rely on the burrows of taxonomic descriptions for kit fox of the Rocky Mountains in the west other species for survival, making use and swift fox. to the Souris hills of in the of them as escape terrain and den In 1978, the Committee On the east, north to the banks of Canada's sites. Swift fox are, in general, more Status of Endangered Wildlife In North Saskatchewan River and south active at night than during the day but Canada (COSEWIC) classified the to the Texas panhandle and the New this activity pattern varies with the swift fox as extirpated in Canada. By Mexico border in the U.S.A season (Pruss 1994; Teeling 1996). 1995, the species was considered ex- (Mackenzie 1911; Carlington 1980; A subspecies of the swift fox, the tirpated from 90% of its historic MacGregor 1998). northern swift fox, V v. hebes, de- range in the USA (United States Fed- The swift fox is North America's scribed by Merriam (1902) was eral Register Vol. 60, No. 116,1995). smallest canid, weighing between 2.3 briefly listed as endangered in the The causes of the extirpation of this and 3.2 kg. They are opportunistic U.S. The species was delisted on the species were attributed to the rapid feeders, eating seeds, berries, grass, basis that a valid subspecific varia- and radical change of the great plains insects, amphibians, reptiles, and tion did not exist, supported by Hall's ecosystem from native grasslands to small animals and birds (Uresk and statement that Vulpes velox was con- cultivated farmland, and the inevi- Sharps 1986; Bremner 1997). If specific with Vulpes macrotis (Hall table hunting, trapping and poison- water is available they require 210 g 1981). By 1980, the northern swift ing programs which accompany such

Vol. 18 No. 4 2001 Endangered Species UPDATE 167 habitat transformation (Carlington projects to be conducted as M.Sc. the- introduction sites appeared suitable, 1980; Weagle and Smeeton 1995). ses (Carlington 1980; Reynolds and offered to provide release sites, Captive breeding of the swift fox 1983; Schroeder 1987) on potential protection and monitoring for the for reintroduction into their original release sites and methodology. The projected swift fox program. Canadian range was initiated in 1972, Canadian Wildlife Service (CWS) a At the Swift Fox Symposium in at the Wildlife Reserve of Western branch of the Federal Government of Saskatoon, Saskatchewan, February Canada (now the Cochrane Ecologi- Canada, became involved for the first 1998 the project was further devel- cal Institute, CEI), using four founder time in the program in 1978, when oped and, as result of these discus- foxes from the United States. Over the Committee on the Status of En- sions, Defenders of Wildlife were the period of the program CEI has dangered Wildlife in Canada contacted to explore project funding. continued to acquire swift foxes to (COSEWIC) designated the swift fox In June 1998, The BFWD invited add to the captive colony from the as "extirpated" in Canada. The Ca- the Directors of CEI to the Blackfeet USA. No Canadian animals were nadian federal government's jurisdic- tribal lands to examine possible swift available for addition to the swift fox tional involvement in the swift fox fox reintroduction sites (Figure 1). captive breeding group as the species reintroduction program was joined by After this meeting and the examination had been extirpated from that coun- provincial government involvement of the potential release sites BFWD try. The captive breeding program when the provinces of Alberta (1983) initiated the project by requesting CEI was initiated solely to provide swift and Saskatchewan (1985) permitted to provide swift fox to the Blackfeet foxes for reintroduction and not for swift fox releases within their juris- Nation for a reintroduction program. exhibit, sale or trade. M.R. Smeeton, diction. The last swift fox reintro- In July of 1998 an 11 day of the CEI, started the swift fox stud- duction took place in Canada in 1997, baseline survey of potential swift fox book in 1972 and in 1986 the stud- and in 1998 the species was habitat, escape terrain, and prey book information was transferred to downlisted from extirpated in Canada abundance and predator abundance ISIS software. That studbook has to endangered in Canada by was conducted to find suitable swift now been consistently maintained for COSEWIC (COSEWIC1998) fox release sites on the AMS Ranch over 29 years. The ISIS software In the summer of 1996 a rancher on the Blackfeet tribal lands. This (ISIS 1989) is used to ensure maxi- from Montana visited the CEI. Upon survey identified seven release sites mum genetic heterozygosity in new his return to the U.S., he contacted to be used for the initial release, that breeding pairs (inbreeding coefficient the Blackfeet Fish &Wildlife Depart- occurred in late August of 1998. <0.05) and, also, that animals of the ment (BFWD), to discuss the possi- same bloodlines are not repeatedly bilities of reintroducing swift fox into Project design reintroduced into the same geo- their original, historic range on the The project was designed as a part- graphic area. In mid-1980, on the Blackfeet Tribal Lands. Swift fox nership of three groups: Blackfeet advice of the newly formed Swift fox were consid- Propagation Committee, swift foxes ered extir- from the CEI captive breeding colony pated from were sent to three zoos (Calgary, the Blackfeet Edmonton, and Moose Jaw) as breed- tribal lands ing and educational exhibit animals. by the 1950s, By 1997, none of the zoos participat- and extir- ing in the breeding program had swift pated from foxes, having either returned their the State of swift fox stock to the CEI, released, Montana by or euthanized them. the 1960s After founding the CEI's captive (Knowles breeding colony in 1972, Miles & 1991). The Beryl Smeeton went on to sign a co- Blackfeet operative agreement with Dr. Steven Tribe showed Herrero, then Dean of the Faculty of interest in the Environmental Design at the Univer- project, pro- sity of Calgary, in 1977. This agree- vided that the Figure 1. Land use in the release areas on the Blackfeet Tribal ment outlined a series of research proposed re- lands, Montana

168 Endangered Species UPDATE Vol. 18 No. 4 2001 Fish and Wildlife Department who Table 1. Summary of data on the swift fox released on Blackfeet provided the lands for the release, Tribal Land, Montana from 1998 to 2000. protection of the reintroduced ani- SwiR Fox Released Cubs Found Recorded Deaths Year Radio Collared mals and some post release monitor- (Fall) (Spring) ing; Cochrane Ecological Institute who maintain the captive colony, and 1998 30 0 NI A 2 provided swift foxes for release, as well as the logistical planning for the 4 releases, including liaison with gov- 1999 8 2 15 (possibly 9) ernment agencies in USA and Canada, development of nonintrusive 3 2000 31 16 2Ot survey methodology and research (to mid October) from graduate students; and Defend- ers of Wildlife who provided a sig- nificant proportion of the funding and ing the release (1998199) the reintro- the cubs sighted on the Blackfeet land help obtaining the requisite additional duced animals were monitored, us- in 2000 appeared to be older than the funding needed for the project. ing tracks and scat, by officers of the cubs born to new pairs at the CEI in The protocol used to release the Blackfeet Fish & Wildlife Depart- 2000. This indicated either: swift foxes on Blackfeet land fol- ment. A spring survey by CEI found (1) that the difference in latitude lowed that described in Smeeton and abundant swift fox sign, three active between the more northerly CEI and Weagle (2000). After identifying dens, one pair with a litter of four Blackfeet lands to the south resulted potential release sites through a pre- cubs and one other den with a pos- in an earlier mating season, or release survey, Portable Protective sible litter of three. A later survey, in (2) that these cubs had been born Shelters (PPS) were set out in the re- early August, discovered a den con- to established pairs, rather than new lease sites. One set of observers and taining 2 cubs. Tho known mortali- pairs, either to the adult pairs released one PPS was used per sibling group. ties occurred during the first winter, in 1999 or juvenile pairs, which had Each sibling group was released in the cause of which was vehicle im- bred in 1999 and again in 2000. Es- the vicinity of the PPS and, for a pe- pact, as the swift foxes were hunting tablished pairs do not go through the riod of two days to one month, the on or near roads. extended mating ritual that newly released animals were kept under In 1999, 15 mixed adult and ju- paired animals do, as a result their cubs close observation. At the end of 10 venile swift foxes were released on are bigger because they are born ear- days the PPS was removed from the the AMS Ranch, eight of these, all lier in the year (Smeeton et al. in press). site. The use of PPS encouraged the adult animals, were radio collared. A In August, 2000 3 1 swift fox animals to remain in the area where radio telemetry program, undertaken were released on the AMS Ranch. they were released. Longterrn moni- by the Blackfeet Tribal Fish &Wild- Sixteen of these animals were radio toring of the population after the first life Department in the spring of 2000, collared. To date there have been month of intensive observation was found 5 of the collared swift fox and three mortalities, two of the mortali- conducted and coordinated by additional searches in August found ties occurred in the first two weeks Blackfeet Fish and Wildlife and the a sixth radio collared swift fox. Al- after the release. CEI. Subsequent swift fox releases though the analysis of the data is not Monitoring of the radio collared took place in areas where pre-release complete first indications are that 75% animals released on Blackfeet tribal surveys indicated swift fox presence. of the captive bred, radio-collared swift lands continues over the winter of foxes survived the winter. 2000 to 200 1. Results to date In 2000, a spring survey under- Table 1 summarizes the swift fox re- taken by the CEI and the Blackfeet Conclusion leased to date on the Blackfeet Tribal Fish and Wildlife Branch discovered The Blackfeet swift fox reintroduc- lands. In 1998,30juvenile swift fox five swift fox natal den sites contain- tion project seems to have benefited were released on the AMS Ranch on ing more than 21 cubs. These cubs from the lessons learned in the Ca- the Blackfeet tribal lands. This was were all born to swift foxes without nadian Swift Fox Reintroduction Pro- the first attempt at reintroducing swift radio collars. gram (1983 to 1997). The use of the fox to be undertaken in the USA. In general, (although they were PPS release methodology, developed Throughout the first winter follow- not trapped and handled to confirm) by the CEI over the latter part of the

Vol. 18 No. 42001 Endangered Species UPDATE 169 Canadian reintroduction program, Green), the U.S. Fish &Wildlife Ser- Merriam, C.H. 1902. Three new foxes of the encouraged the animals to remain in vice, and the State of Montana and kit and desert fox group. Proceedings of the release sites and reduced mortal- the Biological Society of Washington its agencies. The CEI captive breed- 15:73-74. ity. The full support for the program ing colony would not have the wide Pruss, S.D. 1994. An Observational Natal by the Blackfeet people and the pro- genetic heterozygosity which it has Den Study of Wild Swift fox (Vulpes tection provided to the released ani- were it not for the founder foxes con- velox) on the Canadian Prairie. Unpub- mals by the Blackfeet Fish & Wild- tributed to the captive breeding lished M.S. thesis, University of Calgary, Calgary, Alberta, Canada. life Department has ensured that none colony by Ms. Vona Bate, Jon Sharps, Reynolds, J. 1983. A Plan for the Reintro- of the animals released on Blackfeet CWB, and the States of Wyoming duction of the Swift Fox to the Canadian lands, unlike those released in the and Colorado. Above all, the CEI Prairies. Unpublished M.S. thesis, Univer- Canadian program, were killed by would be lost without the work of my sity of Calgary, Calgary, Alberta, Canada. being accidentally poisoned, trapped, Schroeder, C. 1987. Swift Fox Reproductive husband Ken Weagle and my friend Biology in Captivity. Unpublished report, or shot. Pre-release surveys to iden- and collaborator, Sian Waters. University of Calgary, Calgary, AB, tify suitable release sites, undertaken Canada. by the CEI on the Blackfeet Tribal Literature cited Smeeton, C. and K. Weagle. 2000. The re- Lands, indicated that the area was Bremner, S. 1997. Diet and hunting introduction of the swift fox Vulpes velox to South Central Saskatchewan, Canada. suitable and that there was sufficient behaviour of captive bred swift fox (Vulpes velox veloxlVulpes velox hebes) intended Oryx 34:171-179 escape- terrain and a large enough for release. Unpubublished M.S. thesis, Smeeton, C., K.Weagle, and S.S.Waters. In prey base to support reintroduction. University of Edinburgh, Scotland. Press. Captive breeding of the swift fox at The survival and successful repro- Carlington, B.G. 1980. Reintroduction of the the Cochrane Ecological Institute in duction of the swift foxes released in Swift fox (Vulpes velox) to the Canadian Alberta, Canada. In L.C. Carbyn and M.A. Sovada, eds. Ecology and Management Montana on the Blackfeet tribal lands Prairie. Unpublished M.S. thesis, Univer- sity of Calgary, Calgary, Alberta, Canada. of Swift fox in a Changing World. Cana- appears to confirm that assessment. Committee on the Status of Endangered dian Plains Research Centre. Unlike the Canadian reintroduction Wildlife in Canada. 1978. Official Classi- Stromberg, M. R andM.S. Boyse. 1986. Sys- program, only captive bred swift fication of the Swift Fox as Extirpated in tematics and conservation of the swift fox, Vulpes velox, in North America. Biologi- foxes have been reintroduced in the Canada. Vol. 1. COSEWIC Status Reports and Evaluations. Ottawa, Canada. cal Conservation 3597-110. Blackfeet program and these animals Committee on the Status of Endangered Teeling. E. C. 1996. Born to be Free: Cap- have survived, bred, and successfully Wildlife in Canada. 1998. Official Classi- tive Breeding as a Conservation Tech- raised their young on the release sites. fication of the Swift Fox as Extirpated in nique. Unpublished M.S. thesis, Univer- Radio telemetry data has shown 75% Canada. Vol. 1. COSEWIC Status Reports sity of Edinburgh, Scotland. Uresk, D. W. and J. C. Sharps. 1986. Den- survivorship over one year from and Evaluations. Ottawa, Canada. Flaherty, M. and R. Plaake. 1986. Response ning habitat and diet of the swift fox in those radio collared captive bred of swift fox (Vulpes velox) to water stress. western . The Great Basin swift foxes that have been reintro- The Journal of Colorado-Wyoming Acad- Naturalist 46:249-253. duced onto the Blackfeet tribal lands. emy of Science 18:51. US Fish and Wildlife. 1994. Endangered and Hall, E.R. 1981. The Mammals of North Threatened Wildlife and Plants. 5OCFR AmericaVol. 2. John Wiley and Sons. New 17.11 & 17.12, United States Fish and Acknowledgements York. Wildlife Service, Washington D.C., USA. The project could not have been un- ISIS. 1989. Single Population Analysis and US Fish and Wildlife. 1995. Twelve Month dertaken without the support of Record Keeping System. International Administrative Finding on Petition to List Species Information System, 12101 the Swift Fox (Action Needed), From Re- Minette Johnson and Defenders of gional Director, Region 6 to Director Wildlife, the Shared Earth Founda- Johnny Cake Ridge Road, Apple Valley, MN. 55124. (AESITE). United States Fish and Wild- tion, the International Wildlife Coa- Knowles, C. 1991. An Ecological and Taxo- life Service, Washington D.C., USA. lition, Elsa Wild Animal Appeal of nomic Review of the Swift Fox (Vulpes Wayne, R. 1998. A look at swift foxlkit fox Canada and all their funders and as- velox) with Special Reference to Montana. taxonomy within the canid complex. First North American Swift Fox Symposium, sociates. Moving large numbers of an FaunaWest Wildlife Consultants, Boulder, MT. Ecology and Conservation of the Swift endangered species across an inter- Macgregor, J.G. 1998. Peter Fidler, Canada's Fox in a Changing World, Saskatoon, national border requires the help and forgotten explorer 1769 to 1822. House Saskatchewan, Canada. support of many jurisdictions, we Ltd., Calgary, AB, Canada. Weagle, K. and C. Smeeton. 1995. thank Ira Newbreast and the Mackenzie, A. 19 11. Voyages from Montreal Behavioural aspects of the swift fox (Vulpes velox) reintroduction program. Pp Blackfeet Fish & Wildlife Depart- through the Continent of North America to the frozen and Pacific Oceans in 1789 268-286 in B. Holst, ed. Proceedings of ment, the province of Alberta, the and 1793. Vol. 1 andVol. 2. Courier Press the 2nd International Conference on En- government of Canada, the Canadian Ltd. Toronto vironmental Enrichment. Copenhagen Food Inspection Agency (Dr. Tom Zoo, Copenhagen, Denmark.

170 Endangered Species UPDATE Voi. 18 No. 4 2001 lnterspecific Interactions Among Wild Canids: Implications for the Conservation of Endangered San Joaquin Kit Foxes Brian L. Cypher Research Ec'ologist, Endangered Species Recovery Program, P.O. Box 9622,Bakersfield, CA 93389;(661) 837-5061; (fax) (661)398-0549; [email protected] Howard 0. Clark, Jr. Ecologist, Endangered Species Recovery Program, 1900 N. Gateway Blvd., #101,Fresno, CA 93727;(559) 453-1103; (fax) (559)453-1227; hclarkQesrp.org Patrick A. Kelly Director, Endangered Species Recovery Program, 1900 N. Gateway Blvd., #I 01,Fresno, CA 93727;(559) 453-1 103; (fax) (559)453-1227; [email protected] Christine Van Horn Job Ecologist, Endangered Species Recovery Program, 3517 Sedona Way, Bakersfield, CA 93309;(661) 834-6781 ; cvanjobQaol.com Gregory D. Warrick Manager, Center for Natural Lands Management, 10018 Rain Check Drive, Bakersfield, CA 93312;(661) 399-2257; cnlmgret9aol.com Daniel F, Williams Executive Director, Endangered Species Recovery Program, Department of Biological Sciences, California State University - Stanislaus, Turlock, CA 95382;(209) 667-3446; (fax) (209)667-3694; [email protected].

Abstract Interspecific interactions among wild canids have significant implicationsfor the conservation and recovery of endangered Sun Joaquin kit foxes (Vulpes macrotis mutica). Coyotes (Canis latrans) and non-native red foxes (V. vulpes) both engage in interference and exploitative compe- tition with kit foxes. Several behavioral and ecological adaptations of kit foxes ameliorate such competition with coyotes and facilitate their coexistence. These adaptations include habitat partitioning, food partitioning, opportunisticforaging patterns, and year-round use of multiple dens. These adaptations are less effective against red foxes due to greater food and habitat overlap, the ability to pursue kit foxes into dens, and high potential for disease transmission. Thus, non-native red foxes pose a serious threat to kit foxes. Interactions between coyotes and red foxes may benefit kit foxes. In particulal; integerence competition by coyotes may limit the abundance and distribution of red foxes in the Sun Joaquin Valley. These interactions should be considered when evaluating management options (e.g., predator control).

Introduction Fur harvests, predator control pro- erally Endangered in 1967 and Califor- Kit foxes are relatively small canids grams, and rodent control programs nia Threatened in 197 1. (1.7 to 3.0 kg) that occur in arid and also may have contributed to observed Interspecific competition from semi-arid habitats of the southwestern declines. San Joaquin kit foxes persist other mammalian predators is an im- United States and northern Mexico. as a metapopulation comprising three portant factor affecting the remaining The San Joaquin kit fox (Vulpes larger "core" populations and a num- San Joaquin kit fox populations macrotis mutica) is a genetically dis- ber of smaller "satellite" populations. (Cypher et al. in press). Coyotes (Ca- tinct subspecies that historically oc- Current threats include continuing habi- nis latrans), (Lynx ws),gray curred in the San Joaquin, Salinas, and tat conversion,rodenticide use, and in- foxes ( cineareoargenteus), Cuyama Valleys of central California. terspecific competition. Much of the (Taxidea taxus), feral cats The abundance and range of this taxon remaining habitat is fragmented, dis- (Felis domesticus),and non-native red have been sigrdicantly reduced, prirna- turbed, and subject to competing land foxes (V vu2pes) all engage in interfer- rily due to habitat loss and degradation uses such as hydrocarbon production ence andfor exploitative competition associated with agricultural, industrial, and water banking (FWS 1998). The with kit foxes. Interference competi- and urban development (FWS 1998). San Joaquin kit fox was listed as Fed- tion consists of direct mortality, harass-

Vol. 18 No. 4 2001 Endangered Specles UPDATE 171 than predation. Coyotes commonly do abundance in some areas have been not consume the kit foxes they kill associated with increases in coyote (Spiegel and Disney 1996;Cypher and abundance (Cypher et al. 2000; White Spencer 1998), although over half of et al. 2000). Coyote predation, how- kit foxes killed at one location were ever, does not appear to be the primary consumed during a period of low food factor driving kit fox population trends. availability (Ralls and White 1995). Instead, food availability as mediated Another effect of interference by annual environmental conditions competition is spatial exclusion in (particularly precipitation) appears to which the presence of one species in be the primary factor influencing kit fox an area results in decreased use of that population dynamics (White et al. area by another species. On a re- 1996; Cypher et al. 2000). Predation gional scale, kit foxes usually are ab- by coyotes potentially could have a sent or less abundant in more rugged more significant impact on kit fox terrain (areas with average slopes populations when food availability is >5%). Kit foxes may have more dif- low or when kit fox populations are small ficulty eluding predators in rugged (particularly introduced populations). terrain. At one study site, kit foxes Coyotes have historically occurred were abundant in rugged terrain when throughout the range of the kit fox, and San Joaquin kit fox (Vulpes macrotis regional coyote abundance was low, kit foxes have evolved adaptive strate- mutica). Photograph by B. Moose but kit fox numbers quickly declined gies for coexisting with coyotes. One PetersonMIRP. in this terrain as coyote abundance such strategy is year-round use of dens. increased (Warrick and Cypher Kit foxes may have over 100 dens scat- ment, and spatial exclusion. Exploit- 1998). On a local scale, White et al. tered throughout their home range, al- ative competition consists of overlap (1994) did not detect any temporal though on average about 12 dens are in use of potentially limited resources segregation among telemetered kit used by a given fox annually such as food items and dens. Competi- foxes and coyotes, indicating that ar- (Koopman et al. 1998). These dens tive interactions with coyotes and red eas were being used concurrently by facilitate escape from predators (Whtte foxes have the greatest implications for the two species. et al. 1994;Cypher and Spencer 1998). San Joaquin kit foxes. Our objectives With regards to exploitative com- In addition, some amount of resource for this paper are to summarize the petition, coyotes consume some of the partitioning occurs between coyotes competitive interactions that occur be- same food items consumed by kit and kit foxes. Although food habits tween kit foxes, coyotes, and red foxes, foxes. Items commonly used by both overlap, coyotes take higher propor- and to assess the potential implications species include black-tailed hares tions of leporids while kit foxes usu- of these interactions for kit fox conser- (Lcpus californicus), desert cottontails ally consume higher proportions of vation and recovery. (Sylvilagus audubonii), kangaroo rats nocturnal rodents, particularly kanga- (Dipodomys spp.), pocket mice roo rats and pocket mice (White et al. Competitive interactions (Chaetodipus californicus, 1995; Cypher and Spencer 1998). Fur- Coyotes engage in both interference Perognathus inornatus), California thermore, there is some evidence that and exploitative competition with kit ground squirrels (Spermophilus some degree of habitat partitioning may foxes. Coyotes have long been recog- beechyi), pocket gophers (Thommys occur with kit foxes prefening more open nized as a significant cause of mortal- bottae), (Acrididae), axas with reduced shtub cover (White et ity for kit foxes (Seton 1925). At vari- Jerusalem crickets (Gryllacrididae), al. 1995;Warrick and Cy-pher 1998). ous study sites throughout the range of and (Eleodes spp.). Overlap in Red foxes also engage in both in- the San Joaquin kit fox, coyotes are the resource use varies with the relative terference and exploitative competition primary source of kit fox mortality for availability of different food items with kit foxes. Historically, native red which the cause of death is identifiable (Whlte et al. 1995; Cypher and Spen- foxes only occurred at high elevations (Hall 1983;Briden et al. 1992;Standley cer 1998), and therefore may not be a in the Sierra Nevada and Cascade et al. 1992; Ralls and White 1995; siguficant factor in all areas or all years. ranges in California, and were not Spiegel and Disney 1996; Cypher et al. The overall effect of competitive found in any portion of the range of 2000). This mortality indeed appears interactions between coyotes and kit San Joaquin kit foxes (Grinnell et al. to be the result of competition rather foxes is not clear. Declines in kit fox 1937). In more recent times, red foxes

172 Endangered Species UPDATE Vol. 18 No. 4 2001 from the eastern United States have colonizing kit fox habitat lacking in killed by coyotes (Clark 2001). Coy- been introduced into lower elevation nearby water sources. Most otes have been suggested as a biologi- areas of California by humans for hunt- observations in the San Joaquin Valley cal control strategy for red foxes in ing and trapping, and have escaped are in relatively close proximity to coastal areas of California where the from fur farms (Jurek 1992; Lewis et sources of water. foxes are preying on endangered Cali- al. 1993). These highly adaptable, non- Interactions between coyotes and fornia least terns (Sterna antillarum native red foxes have spread rapidly red foxes can affect kit foxes. The pres- browni) and California light-footed and have colonized many regions of ence of both coyotes and red foxes in a clapper rails (Rallus longirostris California, including the San Joaquin given area may act in an additive man- levipes) (Jurek 1992). Coyotes also Valley. The abundance of anthropo- ner with regards to reducing food avail- have been proposed as a means of re- genic water sources in the San Joaquin ability for kit foxes. Thus, exploitation ducing red foxes in the prairie pothole Valley (e.g., canals, irrigated agriculture, competition between coyotes and red region of North America, thereby re- stock ponds, and urban areas) likely has foxes may detrimentally sect kit foxes. ducing red fox predation on duck nests fkilitated colonization by red foxes. (Sargeant and Arnold 1984). Red foxes are larger (3 to 8 kg) than kit foxes, and therefore dominate in conservation implications interference competition. Red foxes are Due to anthropogenic ecosystem known to have killed radiocollared kit modification, non-native red foxes foxes on at least three occasions (Ralls are expanding their range and in- and White 1995; Clark 2001). In addi- creasing in abundance in California tion, red foxes may be competitively (Jurek 1992; Lewis et al. 1993). As displacing kit foxes in some locations described above, this species poses (Whlte et al. 2000). At one location, a potentially serious threat to kit kit foxes primarily used areas not oc- foxes. Implementing effective con- cupied by red foxes (Clark 2001). Fi- trol programs for red foxes would be nally, red foxes and kit foxes are closely extremely difficult for a number of related taxonomically which may increase reasons. In general, predator control the potential for disease transmission. programs can be costly as they usu- Being relatively close in size to kit ally must be implemented over large foxes, exploitative competition may be areas for multiple years (sometimes more intense between the two species. indefinitely) to achieve effective con- Considerable overlap in use of foods trol, and are not popular with the gen- was documented at one location in the eral public even when conducted for San Joaquin Valley (Warrick and Clark the conservation of a rare, native spe- unpublished data). Red foxes will use cies (Goodrich and Buskirk 1'995). kit fox dens, excluding use by kit foxes. Poisons can not be used due to the Red foxes have been observed in dens threat to kit foxes as well as other formerly used by kit foxes on a number non-target species. Trapping red of occasions at three different locations Interference competition between foxes without also capturing kit foxes (white et al. 2000; Cypher et al. in press; coyotes and red foxes may also benefit would be difficult due to the rela- Wanick and Clark unpublished data). kit foxes. Reduced abundance of red tively close weights of the two spe- Red foxes constitute a serious foxes attributable to coyotes has been cies. The use of many types of trap- threat to San Joaquin kit foxes. Adap- documented in a number of locations ping devices was banned in California tations of kit foxes that reduce compe- (Dekker 1983; Voigt and Earle 1983; in 1998. Shooting, possibly in conjunc- tition with coyotes are less effective Major and Sherburne 1987; Sargeant tion with predator calling, may be pos- against red foxes. Red foxes can enter et al. 1987). This reduction is a conse- sible, but could be difficult and costly most dens used by kit foxes, making quence of both direct mortality and to implement over large areas or near escape and avoidance more difficult. exclusion. In a study in the San Joaquin human-inhabited areas where red foxes Resource partitioning may be minimal Valley of California, 11 radiocollared commonly occur. although some degree of habitat parti- red foxes were recovered dead. Of the Coyote control is often suggested tioning may occur between the two nine mortalities for which the cause of as an action that would benefit San species. Red foxes may have difficulty death could be identified, all nine were Joaquin kit foxes. Similar to control

Vol. 18 No. 4 2001 Endangemd Species UPDATE 173 efforts for red foxes, coyote control can ergy Commission, Sacramento. Bird and Mammal Section Report 93-10. be both difficult and controversial Clark, H.O., Jr. 2001. Endangered San Joaquin Major, J. T., and J. A. Sherbwne. 1987. Inter- kit fox and non-native red fox interspecific specific relationships of coyotes, bobcats, (Cypher and Scrivner 1992; Connolly competitive interactions. M.S. thesis, Cali- and red foxes in western Maine. Journal of and Longhurst 1975). There may be fornia State University, Fresno. Wildlife Management 51:606-616. certain situations where limited coyote Connolly, G. E., and W. M. Longhurst. 1975. Ralls, K., and P, J. White. 1995. Predation on control might benefit kit foxes (e.g., re- The effects of control of coyote populations: San Joaquin kit foxes by larger canids. Jour- introduction sites, smaller preserves a simulation model. University of Califor- nal of Mammalogy 76:723-729. nia - Davis, Cooperative Extension Bulle- Sargeant,A. B., S. H. Allen, and J. 0.Hastings. and habitat blocks during periods of tin 1872. 1987. Spatial relations between sympatric low food availability). Given the po- Cypher, B. L., P. A. Kelly, and D. F. Williams. coyotes and red foxes in North Dakota. Jour- tential beneficial role of coyotes in lim- In press. Factors influencing populations of nal of Wildlife Management 51:285-293. iting non-native red foxes, however, the endangered San Joaquin kit foxes: implica- Sargeant, A. B., and P, M. Arnold. 1984. Preda- tions for conservation and recovery. Pro- tor management for ducks on waterfowl pro- implementation of coyote control ceedings of the North American swift fox duction areas in the northern plains. Pp 161 - should be carefully considered. Any symposium. 167 in Proceedings of the 1 lth vertebrate pest reduction or limitation of red fox abun- Cypher, B. L., and J. H. Scrivner. 1992. Coy- control conference. University of Califor- dance achieved naturally through com- ote control to protect endangered San nia, Davis. petitive pressure from native predators Joaquin kit foxes at the Naval Petroleum Seton, E. T. 1925. Lives of game animals. Reserves, California. Pp 42-47 in J. E. Volume I: cats, wolves and foxes. could simcantly benefit kit foxes and Borrecco and R. E. Marsh, eds. Proceed- Doubleday, Doran and Company, New York. would require no effort on the part of ings of the fifteenth vertebrate pest confer- Spiegel, L. K., and M. Disney. 1996. Mortal- humans. Red foxes are rarely observed ence. University of California, Davis. ity sources and survival rates of San Joaquin in areas where coyotes are abundant, Cypher, B. L., and K. A. Spencer. 1998. Com- kit fox in oil-developed and undeveloped petitive interactions between coyotes and lands of southwestern Kern County, Califor- even though kit foxes persist in these San Joaquin kit foxes. Journal of Mammal- nia. 4 71-92 in L. K. Spiegel, ed. Studies areas (Ralls and White 1995; Spiegel ogy 79:204-214. of the San Joaquin kit fox in undeveloped and Disney 1996; Cypher et al. 2000). Cypher, B. L., G. D. Warrick, M. R. M. Otten, and oil-developed areas. California Energy Ultimately, the strategy with the T. P. O'Farrell, W. H. Beny, C. E. Harris, T. Commission, Sacramento. greatest potential for effectively con- T. Kato, P. M. McCue, J. H. Scrivner, and B. Standley, W. G., W. H. Beny, T. P. O'Farrell, W. Zoellick. 2000. Population dynamics of and T. T. Kato. 1992. Mortality of San serving and recovering San Joaquin kit San Joaquin kit foxes at the Naval Petroleum Joaquin kit fox at Camp Roberts Army Na- foxes will be to conserve and properly Reserves in California. Wildlife Mono- tional Guard Training Site, California. U.S. manage large blocks of habitat that are graphs 145. Department of Energy Topical Report No. connected by movement corridors. Dekker, D. 1983. Denning and foraging hab- EGG 10627-2157. its of red foxes, Vulpes vulpes, and their in- U.S. Fish and Wildlife Service. 1998. Recov- This will facilitate larger kit fox popu- teractions with coyotes, Canis latrans, in ery plan for upland species of the San Joaquin lations that are more robust to losses central Alberta, 1972-198 1. Canadian Field- Valley, California. U.S. Fish and Wildlife from interspecific competition, and that Naturalist 97:303-306. Service, Portland, Oregon. are able to naturally recolonize areas Goodrich, J. M., and S. W. Buskirk. 1995. Voigt, D. R., and B. D. Earle. 1983. Avoid- where local extirpations of kit foxes Control of abundant native vertebrates for ance of coyotes by red fox families. Journal conservation of endangered species. Con- of Wildlife Management 471852-857. may have occurred. Also, the fragmen- servation Biology 9: 1357-1364. Warrick;G. D., and B. L. Cypher. 1998. Fac- tation of kit fox habitat by anthropo- Grinnell, J., J. S. Dixon, and J. M. Linsdale. 1937. tors affecting the spatial distribution of a kit genic disturbances promotes increased Fur-bearing mammals of California, Vol. 2. fox population. Journal of Wildlife Man- abundance of red foxes, which nega- University of California Press, Berkeley. agement 62:707-717. Hall, F, A., Jr. 1983. Status of the San Joaquin White, P, J., W. H. Berry, J. J. Eliason, and M. tively impacts kit foxes. The conser- kit fox, Vulpes macrotis mutica, at the T. Hanson. 2000. Catastrophic decrease in vation of large blocks of habitat is a Bethany wind turbine generating (WTG) an isolated population of kit foxes. South- paramount goal of the recovery plan for project site, Alameda County, California. western Naturalist 45204-21 1. San Joaquin kit foxes and a number of California Department of Fish and Game, White, P,J., K. Ralls, and R. A. Garrott. 1994. other rare species that occur sympatri- Sacramento. Coyote-kit fox interactions as revealed by Jurek, R. M. 1992. Nonnative red foxes in telemetry. Canadian Journal of zoology cally with kit foxes (FWS 1998). California. California Department of Fish 72:1831-1836. and Game, Nongame Bird and Mammal White, P. J., K. Ralls, and C. A. Vanderbilt Literature cited Section Report 92-04. White. 1995. Overlap in habitat and food Briden, L.E., M. Archon, and D.L. Chesemore. Koopman, M. E., J. H. Scrivner, and T.T. Kato. use between coyotes and San Joaquin kit 1992. Ecology of the San Joaquin kit fox in 1998. Patterns of den use by San Joaquin foxes. Southwestern Naturalist 40:342-349. Western Merced County, California. Pp 8 1-87 kit foxes. Journal of Wildlife Management White, P. J., C. A. Vanderbilt White, and K. in D. E Warn, S. Byme, and T. A. Rado, 62:373-379. Ralls. 1996. Functional and numerical re- eds. Endangered and sensitive species of the Lewis, J. C., K. L. Sallee, and R. T.Golightly, Jr. sponses of kit foxes to a short-term decline San Joaquin Valley, California: their biology, 1993. Introduced red fox in California Cali- in mammalian prey. Journal of Mamrnal- management, and conservation. Califomia En- fornia Department of Fish and Game,Nongame ogy 77:370-376.

174 Endangered Species UPDATE Vol. 78 No. 4 2001 Predation Management Chemical Repellents and Other Aversive Strategies in Predation Management J.R. Mason National Wildlife Research Center, USDA-APHIS-WS, BNR-163, Utah State University, Logan, UT 84322-5295; (435) 245-6091; James.R.Mason Q usda.gov J.A. Shivik National Wildlife Research Center, USDA-APHIS-WS, 4101 LaPorte Avenue, Fort Collins, CO 80521-2154; (970) 266- 6088; John.ShivikQusda.gov M.W. Fall National Wildlife Research Center, USDA-APHIS-WS, 4101 LaPorte Avenue, Fort Collins, CO 80521-21 54; (970) 266- 6086; Michael.W.Fall Q usda.gov

Abstract Chemical repellents and other aversive strategies are the core of non-lethal wildlife management. These strategies typically depend on irritation (pain),conditioning, or fear for their effectiveness, and none is universally successful. Thus, conditioned food aversions deter browsing and forag- ing by deer (Odocoileus virginianus, 0. hemionus), but are less useful with predators, because killing, not consumption, is the behavior of interest. Broadly speaking, the utility of non-lethal strategies is afSected by number and density of wildlife species, availability of alternative foods, palatability and novelty of treated items, and intensity of pain, sickness, or fear used to establish avoidance. Some of the most promising areas for successfil predation management are those involving a combination of strategies tailored to a spec@ problem. For example, behavioral- contingent auditory and visual stimuli coupled with presentations of electric shock or momentary vibration (via telemetry collars) could provide an effective and unambiguous cue for withdrawal. Non-lethal methods, however; are rarely stand-alone technologies. More ofen, integrated strategies, involving both lethal and non-lethal methods, are required for effective predation management.

Introduction creasing despite the fact that effec- ditory cues, chemical and color The survival or restoration of tive options remain virtually non- cues) whenever non-lethal strate- threatened and endangered species existent. Repellents and other aver- gits are implemented. can depend on protection from sive techniques provide cases in predators (Witmer and Fall 1995; point. If wildlife numbers are suf- Chemical repellents Witmer et al. 1996; Hecht and ficiently high, or alternative foods Vertebrate chemical repellents are Nickerson 1999). Most of the rel- are sufficiently scarce, repellents effective because they are irritating, evant data for managing predation usually fail as a deterrent. Few de- cause sickness, or stimulate fear (Ma- stem from research on the protec- monstrably effective alternatives son and Clark 1997). As a rule, these tion of livestock, crops, and com- exist, and practical obstacles to the substances are most useful when they modities (Campbell et. a1 1998; Fall development of new materials are are applied directly to inert materials and Jackson 1998). Deterring considerable. The present discus- (e.g., prepared foods, fruits, grains, predators from prey is even more sion will cover these topics by con- electrical wiring, irrigation hose; complex than protecting crops or sidering: mechanisms underlying Werner et al. 1998). There is no good other commodities because more is the effectiveness of repellents and evidence that predators or other wild- involved than food consumption aversive agents, regulatory con- life will avoid areas protected solely (Fall 1990; Knowlton et al. 1999). straints that govern implementation with border treatments. To illustrate Especially challenging is the devel- of new methods, and the impor- the point, Renardine is commercially opment of non-lethal approaches. tance of employing multiple sen- available for use with red fox (Vulpes Demand for these strategies is in- sory modalities (i.e. visual and au- vulpes) in the United Kingdom and

Vol. 18 No. 4 2001 Endangered Species UPDATE 175 is being evaluated for use with coy- particular coyotes, are markedly in- when prey are completely infused. otes (Canis latrans) in Canada (Mar- sensitive to denatonium benzoate This strategy is relatively common tin and O'Brien 2000). (Mention of (Mason and McConnell 1997). De- among insects (e.g., Wickler 1968), trade names and manufacturers is for spite this lack of demonstrated util- amphibians, reptiles (e.g., Schmidt et identification only and does not im- ity, new veterinary and wildlife con- al. 1989), and occasionally birds. , ply endorsement by the authors or the trol products containing denatonium One example is the Pitohui bird U.S. Department of Agriculture.) derivatives as the active ingredient (Pitohui dichrous) that stores and The substance is bone tar oil dis- are regularly offered for sale. uses toxicants from insects it ingests solved in kerosene. The label ad- (Dumbacher et al. 1992). However, vises that it should be applied lib- Irritation in the absence of complete infusion, erally to pasture borders (on fence Among the three types of chemical repellency is easily circumvented. posts, etc.) to prevent predators repellents, substances that cause sen- This explains the ineffectiveness of from entering and attacking live- sory irritation or pain (the same neu- topically applied irritants as deter- stock. In testing with captive coy- ral receptors are involved) usually rents to predation. For example, otes in the U.S., not only did are most effective. This is because sheep fitted with collars containing Renardine fail to prevent entries sensory pain leads to immediate capsicum oleo resin were killed as into areas, but food adulterated with withdrawal, independent of learn- readily as sheep without collars, de- the material was eaten as rapidly as ing. Such avoidance does not ha- spite the fact that attacking coyotes unadulterated food (Zemlicka and bituate (diminish) for as long as the punctured collars and were exposed Mason 2000). This probably re- irritating stimulus is present. More- to high concentrations of the irritant flects the fact that sulfurous com- over, taxonomic differences in irri- (Burns and Mason 1996). pounds in bone tar oil are attractive tant sensitivity between birds and to coyotes (see Fear below). mammals (Clark 1998a) permit the Fear Tastes, per se, are rarely effec- development of repellents with Sulfur compounds and volatile am- tive feeding deterrents. While bit- some degree of selectivity (e.g., monium soaps of higher fatty acids ter and acidic substances can ini- Norman et al. 1992). induce what humans describe as tially reduce the consumption of For mammals, strong irritants fear in herbivores (Milunas et al. treated materials slightly (cf. Nolte include capsaicin and capsicum oleo 1994). These substances underlie et al.l994b), intake typically re- resins (i.e. the active ingredients in the effectiveness of predator urines turns to baseline within a short pe- 'hot sauce' preparations; Norman et and many commercial preparations riod of time. Products that claim al. 1992), volatile chemicals like used to repel browsing deer, effectiveness solely because of a mustard oil (ally1 isothiocyanate) and (Sylvilagusfloridanus), and rodents "bad" taste are doing so largely be- ammonia (Budavari et al. 1996), and (Nolte et al. 1994a; Lewison et al. cause humans find the taste repul- non-volatile substances including as- 1995; Mason et al. 1999). Typi- sive. Some species of interest, in- tringent tannins such as quebracho cally, substances that frighten her- cluding obligate carnivores such as (Swihart 1990). None of these sub- bivores attract obligate carnivores the Felidae, have taste sensitivities stances repel birds (Mason and Clark and many (Kimball et al. that are greatly different than hu- 1997). Unfortunately, when irritant 2000). There are no published data mans, including insensitivity to salt chemicals dissipate (e.g, by evapo- consistent with the belief that urine and sweet (Beauchamp and Mason ration or photolysis), there is usually samples from one predator are ac- 1991). In particular, products that an immediate resumption of the un- tively avoided by other predators. contain denatonium derivatives wanted behavior (Mason et al. 1985). A disadvantage of fear-inducing (compounds very bitter to humans) A more important drawback is that chemicals is that animals readily ha- are ineffective repellents, almost re- intrataxonornic differences in irritant bituate to their presence. The rate of gardless of species (although bears, sensitivity are small. There are no habituation is largely dependent on Ursus horribilis and Ursus known irritants that affect only some the degree to which the chemical cue americanus, may avoid denatonium mammalian species but not others is associated with a risk of predation. in foods; G. Witmer, National Wild- (e.g., coyotes but not sheep, Ovis When risk is low, habituation is rapid. life Research Center, personal com- aries, or humans). Cues may even become attractive. munication) or method of applica- This is not to say that irritants are For example, wolf (Canis lupis) urine tion (e.g., topical spray, incorpo- completely ineffective deterrents to applied as a repellent along roadways rated into products). Canids, in predation. Irritants can be effective in winter can attract moose (Alces

176 Endangered Species UPDATE Vol. 18 No. 4 2001 alces) and other ungulates that learn tions, no lithium chloride- or estro- consumption of prey, the generali- to associate the odor with the pres- gen-based method has been regis- zation of learning from consump- ence of road salt (T. Sullivan, per- tered by the U.S. Environmental tion to killing is weak. There are sonal communication). Protection Agency. no data on the use of CA to protect Gustavson (1974) conducted big game from predators, but con- Sickness (conditioned or learned the first studies of CA as a manage- siderable data relating to attempts avoidance) ment strategy with coyotes. His ini- at livestock protection provide little When the inge'stion of novel flavors tial data were promising, generat- promise of potential utility. or tastes by mammals or distinctively ing considerable interest in the ap- colored foods by birds is followed by proach. Some investigators have Mechanical and electronic sickness, a learned avoidance usually reported success in preventing pre- devices results (Beauchamp and Mason dation (Gustavson et a1.1974; Ellins There are many parallels between 1991). This effect is variously called and Martin 1981; Gustavson et chemical repellents and mechanical conditioned (or learned) taste, food, a1.1982; Forthman-Quick et al. or electronic devices that provoke or flavor avoidance (CA). CA can 1985a, 1985b), while others have fear or deliver painful or irritating occur after a single aversive experi- reported failure (Conover et al. stimulation. Scarecrows and their ence, particularly when the intensity 1977; Burns 1980; Burns and modern analogues have been of sickness is great and the taste, Connolly 1980; Bourne and widely examined with both birds food, or flavor is new (Pelchat et al. Dorrance 1982; Burns 1983). Two and mammals. Studies typically 1983). As with other chemical repel- large field trials conducted in report rapid habituation and vari- lents, substances that elicit CA are Canada generated opposite results ability among species and settings classified as pesticides by regulatory (Bourne and Dorrance 1982; (Koehler et al. 1990; Bomford and agencies, which typically require ex- Gustavson et al. 1982). Ten years O1Brien 1990). Nevertheless, at tensive data sets for registration prior after the most extensive field trial least for depredation management, to commercial use. (Gustavson et al. 1982; Jelinski et there are promising results in cer- An extensive literature on theory, al. 1983), survey responses of 52 tain situations. For example, use and applications of CA is avail- participating ranchers indicated that Linhart et al. (1984) found that able (e.g., Riley and Tuck 1985). CA while 54% initially considered combinations of battery-operated is the mechanism underlying the util- lithium-chloride baiting "success- strobe lights, sirens, and high fre- ity of commercial bird repellents con- ful" or "somewhat successful," only quency horns, placed on the edges taining methiocarb or anthraquinone one participant continued to use it of sheep pastures or bedgrounds (Conover 1982; Reidinger and Ma- (Conover and Kessler 1994). While and activated for short irregular in- son 1983), and commercial deer, rab- no explanation for differences tervals during night and early morn- bit, and rodent repellents containing among studies is completely ac- ing, stopped predation for 27 to 136 thiram or ziram (Thomson 1995). CA cepted, most arguments have fo- nights. Coyotes apparently re- using lithium chloride or estrogens cused on methods and experimen- mained active around the peripher- to induce sickness has been investi- tal design (Bekoff 1975; Gustavson ies of the test pastures, but habitu- gated as a way to: reduce depreda- et al. 1975; Sterner and Shumake ation to the devices was delayed by tion by coyotes, resolve nuisance 1978; Horn 1983; Forthman-Quick the irregular patterns of activation. feeding by black bears (Ursus et a1.1985b; Conover 1997). Al- More recently, using animal-acti- americanus; Ternent and Garshelis though CA may be a useful tool in vated or demand-performance 1999), and curtail egg predation by some situations, its utility in pre- frightening devices (Stevens et al. raccoons (Procyon lotor), dation management appears quite 2000), Shivik and Martin (in press) ( mephitis), limited. Gustavson acknowledged showed that motion-activated si- ( nyula), and ravens and that coyotes often resumed killing rens were more effective than ran- crows (Corvus spp.; e.g., Nicolaus sheep shortly after conditioning, dom sirens in delaying habituation and Nellis 1987; Nicolaus et al. and ". . .once a coyote becomes a by captive coyotes. Limited field 1982, 1983; Semel and Nicolaus confirmed sheep-killer, perhaps it trials of behavior-contingent, multi- 1992). While evidence suggests will be necessary to remove it from stimulus (light and sound), systems that CA can be used to successfully the population" (Gustavson et al. to deter wolf predation are ongo- manage nuisance complaints and 1978). This could reflect the pos- ing and appear promising (Shivik egg depredation under some condi- sibility that while CA may affect and Martin in press). Devices are

Vol. I8 No. 4 2001 Endangered Specles UPDATE 177 activated when radio-collared wolves target species) often associated with ment station where non-lethal approach livestock production areas. pest control efforts. Much attention methods and non-selective coyote Application of a brief, non-le- has focused in the past on unin- removal had not achieved desired thal electric current has been widely tended consequences on non-target reductions in predation after several studied as a means of deterring species of broad-spectrum pesticide years of effort. Sacks et al. (1999) predators (Linhart et al. 1982; use. However, all proposed meth- found that adult territorial coyotes Sargeant and Arnold 1984). Linhart ods of pest control, including those responsible for sheep killing in the et al. (1976) found that coyotes fit- presumed to be non-lethal, must be area were less vulnerable to these ted with collars that provided a con- carefully examined for effective- control tactics than coyotes not in- tingent electric shock stopped at- ness in specific situations, selectiv- volved in livestock predation. U1- tacks on rabbits for several months. ity, and potential environmental and timately, removal of specific dep- More recently, commercial elec- behavioral effects. For example, redating individuals by shooting tronic dog training collars that de- fences placed to exclude a preda- and Livestock Protection Collars liver a mild static electrical dis- tory species may interrupt move- greatly reduced predation (Blejwas charge have been successfully ment patterns or block migration et al, in press). Livestock Protec- tested as a deterrent to captive coy- routes of another. Selectivity for tion Collars, however, have re- otes attacking sheep (Andelt et al. particular problem species, or indi- cently been banned in California by 1999). Manual activation of col- vidual animals causing predation, a ballot initiative, creating a need lars stopped attacks in progress and and avoidance of problems with for development of effective alter- greatly reduced the probability of primary or secondary effects on native methods. In another situa- subsequent attacks. After one to non-target animals are desirable tion, probably common in both ag- three training bouts, coyotes features for all animal control tac- riculture and conservation preda- avoided or retreated from sheep in tics, especially those involving tion management applications, fe- tests four months after initial ses- chemical applications. Classic ex- ral cat (Felis catus) control efforts sions. Shivik and Martin (in press) amples of past successes in agricul- had to be implemented following a are testing similar collars, triggered ture in finding such alternative ap- highly successful rodent control ef- by radio signals, with wolves. Ani- proaches include: (1) replacing dy- fort to protect nesting Dark-rumped mals wearing modified radio telem- namiting of vampire bat (Diphylla petrels (Pterodroma phaeopygia) etry collars self-activate the static ecaudata) caves to control paralytic from predation in the Galapagos Is- discharge when they approach cattle rabies with vampire bat-se- lands (Cruz and Cruz 1987). When within biting distance of a calf, pro- lective toxicant treatments black rats (Rattus rattus) were re- viding an unambiguous cue for (Mitchell 1986; Lewin 1986); and moved as the primary predator, withdrawal. Collars utilizing mo- (2) replacing poisoned carcass bait cats, which had been subsisting on mentary vibration (a sensation hu- stations as a method of coyote pre- rats, switched prey, diminishing ini- mans perceive as similar to static dation control with selective meth- tial increases in nesting success discharge) are also commercially ods, such as Livestock Protection achieved by the rat control pro- available for dog training and could Collars (Connolly et al. 1978; gram. Although non-lethal tactics, have similar application. Shivik Connolly 1993), den hunting (Till if they become available, would be and Martin (in press) describe ef- and Knowlton 1983), and aerial expected to be more benign and forts to develop auto-attaching col- hunting (Connolly and O'Gara specific in such situations, both le- laring systems that utilize break- 1988; Wagner and Conover 1999; gal and ethical considerations re- away snare technology, which, if and Mason et al. in press). quire their careful assessment be- successful, would substantially re- Similarly, control programs fore implementation takes place. duce the cost of the method. aimed at protecting endangered species from predation must be Regulatory constraints Ecological and behavioral planned strategically for specific Obstacles to development of new consequences areas to assure they achieve desired materials and methods .are consider- At issue is effective adaptation of objectives. For example, Conner able. These include a variety of con- agricultural methods to endangered et al. (1998) found no relationship straints imposed by regulatory agen- species protection, while avoiding between annual coyote removal and cies. Even when new repellent tech- negative ecological consequences levels of coyote predation on sheep nologies are uncovered, the path to (e.g., affecting wildlife other than on a California agricultural experi- commercial availability is long and

178 Endangered Species UPDATE Vol. 18 No. 4 2001 can be very expensive (Fagerstone pulegone (Mason 1990) are both Hillsdale, New Jersey. and Schafer 1998). For example, broadly effective vertebrate (bird and Bekoff, M. 1975. Predation and aversive conditioning in coyotes. Science methyl anthranilate is one of two new mammal) repellents that have both 187: 1096. bird repellents to become commer- irritant and sickness-inducing effects. Blejwas, K.M., Sacks, B.N., Jaeger, M.M., cially available in the United States and D.R. McCullough. In press. The ef- during the past 25 years. This natu- Conclusion fectiveness of selective removal of breed- ral substance is GRAS-listed (gener- Development and application of eco- ing coyotes in reducing sheep depredation. Journal of Wildlife Management. ally recognized as safe) with the U.S. logically sound and effective repel- Bomford, M. and P.H. O'Brien. 1990. Sonic Food and Drug Administration and lents is dependent upon a knowledge deterrents in animal damage control: a re- it has been widely used as a grape of the sensory Umwelt of the species view of device tests and effectiveness. flavoring in human and animal feeds in question (von Uexkull 1934). Wildlife Society Bulletin 18:411-422. Bourne, J., and M.J. Dorrance. 1982. A field since the turn of the century. Despite Even when aversive strategies can be test of lithium chloride aversion to reduce these facts, registration efforts to per- successfully applied, their continued coyote predation on domestic sheep. Jour- mit spraying of methyl anthranilate utility will likely depend on applica- nal of Wildlife Management 46:235-239. on turf to deter grazing Canada geese tion of other techniques in a mosaic Budavari, S., M.J. O'Neil, A. Smith, P.E. (Branta canadensis) took five years of management strategies designed to Heckelman, and J.F. Kinneary, eds. 1996. The Merck Index, 12" Edition. Merck and and cost $5 million (P.J. Voigt, R. J. meet requirements of a specific lo- Company, Whitehouse Station, NJ. Advantage, Inc., personal communi- cation, time, and context. Indeed, se- Burns, R.J. 1983. Microencapsulated cation). While research on non-le- lective removal of wildlife (either in lithium chloride bait aversion did not stop thal methods for agricultural appli- terms of local population suppression coyote predation on sheep. Journal of Wildlife Management 47: 1010- 1017. cations has been relatively well-sup- or removal of specific individuals) Burns, R.J., and G.E. Connolly. 1980. ported, support needed to develop may often be prerequisite for effec- Lithium chloride bait aversion did not in- and evaluate methods for endangered tive implementation of non-lethal al- fluence prey killing by coyotes. Proceed- species applications has been more ternatives. For this reason, integrated ings of the Vertebrate Pest Conference elusive, usually coming in the form strategies that incorporate both lethal 9:200-204. Burns, R.J. and J.R. Mason. 1996. Effec- of small grants that cannot cover long and non-lethal methods will often be tiveness of Vichos non-lethal collars in de- term costs of developmental research the most logical course for effective temng coyote attacks on sheep. Proceed- to meet regulatory requirements. predation management. The high ings of the Vertebrate Pest Conference cost of development and application 17~204-206. Campbell, E.W., M.W. Fall, R.T. Sugihara, Combinations of stimuli of alternative technologies for endan- andA.E. Koehler. 1998. A review of wild- Beauchamp (1997), among others, gered species applications and the life damage management techniques de- has suggested that the most effective highly specific minor-use markets for veloped for use in agriculture that could strategy in development of repellents such products, which limit private in- be used for endangered species conserva- may be use of combinations of dustry interest in the problem, present tion (abstract). Hawaii Conservation Con- ference, Honolulu, HI. stimuli. The evidence is consistent challenges to the emergence of new Clark, L. 1998a. Physiological, ecological, with this suggestion (Clark 1998b). technologies needed to help assure and evolutionary bases for the avoidance Thus, a mixture of capsaicin (irrita- effective recovery of species threat- of chemical initants by birds. Current Or- tion), thiram (sickness-based condi- ened by predation. nithology 14:1-33. tioned avoidance), and Big Game Clark, L. 1998b. Bird repellents: interaction of chemical agents in mixture. Physiol- Repellent (sulfur-based fear) is a Literature cited ogy and Behavior 64:689-695. more effective deterrent to browsing Andelt, W.E, R.L. Phillips, K.S. Gruver, and Conner, M.M., M.M. Jaeger, T.J. Weller, and white-tailed deer (Odocoileus J.W. Guthrie. 1999. Coyote predation on D.R. McCullough. 1998. Effect of coy- virginianus) than any of these sub- domestic sheep deterred with electronic ote removal on sheep predation in north- dog-training collar. Wildlife Society Bul- em California. Journal of Wildlife Man- stances alone (M. Richmond, U.S. letin 27:12-18. agement 62(2):690-699. Geological Survey, Cornell Univer- Beauchamp, G.K. 1997. Chemical signals and Connolly, G. 1993. Livestock protection sity, personnel communication). repellency: problems and prognosis. Pp. 1- collars in the United States, 1988-1993. Likewise, mixtures of methiocarb 10,Pp. 371-392, in J.R. Mason, ed. Repel- Proceedings Great Plains Wildlife Dam- lents in Wildlife Management. Colorado (sickness-based conditioned avoid- age Control Workshop 11:25-33. State University Press, Ft. Collins, CO. Connolly, G.E., R.E. Griffiths, Jr., and P.J. ance) and methyl anthranilate (irri- Beauchamp, G.K. and J.R. Mason. 1991. Savarie. 1978. Toxic collar for control of tation) are more effective than either Comparative hedonics of taste. Pp. 159- sheep-killing coyotes: a progress report. substance alone. Cinnamaldehyde 184 in R.C. Bolles, ed. The Hedonics of Proceedings of the Vertebrate Pest Con- (Crocker and Perry 1990) and d- Taste. Lawrence Erlbaum and Associates. ference 8:197-205.

Vol. 18 No. 4 2001 Endangered Species UPDATE 179 Connolly, G. andB.W. OIGara. 1988. Aerial the coyote and . Ph.D. dissertation. Linhart, S.B., J.D. Roberts, S.A. Shumake, hunting takes sheep-killing coyotes in University of Utah. and ReJohnson. 1976. Avoidance of prey western Montana. Proceedings Great Gustavson, C.R., L.P. Brett, J. Garcia, and by captive coyotes punished by electric Plains Wildlife Damage Control Workshop D.J. Kelly. 1978. A working model and shock. Proceedngs of the Vertebrate Pest 8:184-188. experimental solutions to the control of Conference :302-306. Conover, M.R. 1982. Behavioral techniques predatory behavior. Pp.21-66 in H. Linhart, S.B., R.T. Sterner, G.J. Dasch, and to reduce bird damage to blueberries: Markowitz and V.J. Stevens, eds. Behav- J.W. Theade. 1984. Efficacy of light and methiocarb and hawk-kite predator mod- ior of Captive Wild Animals. Nelson and sound stimuli for reducing coyote preda- els. Wildlife Society Bulletin 10:211-216. Hall, Chicago. 314 pp. tion upon pastured sheep. Protection Ecol- Conover, M.R. 1997. Behavioral principles Gustavson, C.R., J.R. Jowsey, and D.N. ogy 6:75-84. governing conditioned food aversions Milligan. 1982. A 3-year evaluation of Martin, J. and A.OIBrien. 2000. The use based on deceptions. Pp. 29-41 in J. R. taste aversion coyote control in of bone oil (Renardine) as a coyote re- Mason, ed. Repellents in Wildlife Man- Saskatchewan. Journal of Range Manage- pellent on sheep farms in Ontario. Pro- agement, Colorado State University Press, ment 3557-59. ceedings of the Vertebrate Pest Confer- Ft, Collins, CO. Gustavson, C.R., J. Garcia, W.G. Hankins, ence 19:310-314. Conover, M.R., J.G. Francik, and D.E. Miller. and K.W. Rusiniak. 1974. Coyote preda- Mason, J.R. 1990. Evaluation of pulegone 1977. An experimental evaluation of us- tion control by aversive conditioning. as an avian repellent. Journal of Wildlife ing taste aversion to control sheep loss due Science 184581-583. Management 54: 130- 135. to coyote predation. Journal of Wildlife Gustavson, C.R., D.J. Kelly, and J. Garcia. Mason, J.R., W.C. Pitt, andM.J. Bodenchuk. Management 41:775-779. 1975. Predation and aversive condition- In press. Factors influencing the efficiency Conover, M.R. and K.K. Kessler. 1994. Di- ing in coyotes. Science 187: 1096. of fixed-wing aerial hunting for coyotes minished producer participation in an aver- Hecht, A. and P.R. Nickerson. 1999. The in the western United States. International sive conditioning program to reduce coy- need for predator management in conser- Biodeterioration & Biodegradation. ote depredation on sheep. Wildlife Soci- vation of some vulnerable species. En- Mason, J.R. and L. Clark. 1997. Avian re- ety Bulletin 22:229-233. dangered Species Update 16:114-1 18. pellents: options, modes of action, and Crocker, D.R. and S.M. Perry. 1990. Plant Horn, S.W. 1983. An evaluation of preda- economic considerations. Pp. 371-392 in chemistry and bird repellents. Ibis tory suppression in coyotes using lithium J. R. Mason, ed. Repellents in Wildlife 132:300-308. chloride-induced illness. Journal of Wild- Management. Colorado State University Cruz, F. and J.B. Cruz. 1987. Control of life Management 47:999-1009. Press, Ft. Collins, CO. black rats (Rattus rattus) and its effect on Jelinski, D.E., R.C. Rounds, and J.R. Jowsey. Mason, J.R., J.F. Glahn, R.F. Reidinger, and nesting dark-rumped petrels in the 1983. Coyote predation on sheep, and R.A. Dolbeer. 1985. Field trial of dim- Galapagos Islands. Vida Silvestre Neo- control by aversive conditioning in ethyl anthranilate as a nontoxic bird re- tropical l(2): 3-13. Saskatchewan. Journal of Range Manage- pellent in cattle and swine feedlots. Jour- Dumbacher, J.P., B.M. Beehler, T.F. Spande, ment 3616-19. nal of Wildlife Management 49:636-642. H.M. Garraffo, and J.W. Daly. 1992. Koehler, A.E., R.E. Marsh, and T.P. Salmon. Mason, J.R., J. Hollick, B.A. Kimball, and Homobatrachotoxin in the genus Pitohui: 1990. Frightening methods and devices/ J.J. Johnston. 1999. Repellency of Deer Chemical defense in birds? Science stimuli to prevent mammal damage - a re- Away big game repellent to eastern cot- 258:799-80 1. view. Proceedings of the Vertebrate Pest tontail rabbits. Journal of Wildlife Man- Ellins, S.R. and G.C. Martin. 1981. Olfac- Conference 14:168- 173. agement 63:309-314. tory discrimination of lithium chloride by Kimball, B.A., J.R. Mason, F.S. Blom, J.J. Mason, J.R. and J.A. McConnell. 1997. He- the coyote (Canis latrans). Behavioral and Johnston, and D.E. Zemlicka. 2000. De- donic responsiveness of coyotes to 15 Neural Biology 31:214-224. velopment and testing of seven new syn- aqueous taste solutions. Journal of Wild- Fagerstone, K.A. and E.W. Schafer, Jr. 1998. thetic coyote attractants. Journal of Agri- life Research 2:21-24. Status of APHIS vertebrate pesticides and cultural and Food Chemistry 48:1892- Milunas, M.C., A.F. Rhoads and J.R. Mason. drugs. Proceedings of the Vertebrate Pest 1897. 1994. Effectiveness of odor repellents for Conference 18:319-324. Knowlton, F., E.M. Gese, and M.M. Jaeger. protecting ornamental shrubs from brows- Fall, M.W. 1990. Control of coyote preda- 1999. Coyote depredation control: an in- ing by white-tailed deer. Crop Protection tion on livestock- progress in research and terface between biology and management. 13:393-397. development. Proceedings of the Verte- Journal of Range Management (5): Mitchell, G.C. 1986. Vampire bat control brate Pest Conference 14:245-25 1. 398-412. in Latin America. Pp. 152- 164 in National Fall, M.W. and W.B. Jackson. 1998. A new Lewin, R. 1986. Why dynamiting vampire Research Council. Ecological Knowledge era of vertebrate pest control? Interna- bats is wrong. Science 232:24-26. and Environmental Problem-Solving. Na- tional Biodeterioration & Biodegradation Lewison, R., N.J. Bean, E.V. Aronov, and J.R. tional Academy Press, Washington, D.C. 42:85-91. Mason. 1995. Similarities between big Nicolaus, L.K., J.F. Cassel, B.B. Carlson, and Forthman-Quick, D.L., C.R. Gustavson, and game repellent and predator urine repel- C.R. Gustavson. 1983. Taste-aversion K.W. Rusiniak. 1985a. Coyote control lency to white-tailed deer: the importance conditioning of crows to control predation and taste aversion. Appetite 6:253-264. of sulfur and fatty acids. Proceedings of on eggs. Science 220:212-214.Nicolaus, Forthman-Quick, D.L., C.R. Gustavson, and the Eastern Wildlife Damage Control Con- L.K., T.E. Hoffman, and C.R. Gustavson. K.W. Rusiniak. 1985b. Coyotes and taste ference : 145-148. 1982. Taste aversion conditioning in free- aversion: the authors' reply. Appetite Linhart, S.B., J.D. Roberts, and G.J. Dasch. ranging raccoons, Procyon lotor. North- 6:284-290. 1982. Electric fencing reduces coyote pre- west Science 56:165-169. Gustavson, C.R. 1974. Taste aversion con- dation on pastured sheep. Journal of Nicolaus, L.K. and D. Nellis. 1987. The ditioning as a predator control method in Range Management 35276-281. first evaluation of the use of conditioned

180 Endangered Species UPDATE Vol. I8 No. 4 2001 taste aversion to control predation by mon- California ranch. Journal of Wildlife alleviating coyote depredations upon do- gooses upon eggs. Applied Animal Management 63(3):939-949. mestic sheep. Journal of Wildlife Man- Behaviour Science 17:329-334. Sargeant, A.B., and P.M. Arnold. 1984. agement 47(4): 1018- 1025. Nolte D.L., J.R. Mason, G. Epple, E. Aronov, Predator management for ducks on water- Thomson, W.T. 1995. Agricultural Chemi- and D.L. Campbell. 1994a. Why are fowl production areas in the northern cals, Book III: Fumigants, Growth Regu- predator urines aversive to prey? Journal plains. Proceedings of the Vertebrate Pest lators, Repellents, and Rodenticides. of Chemical Ecology 20: 1505-1516. Conference 11:161-167. Thomson Publications, Fresno, CA. Nolte, D.L., J.R. Mason and L. Lewis. Schmidt, P.J., W.C. Sherbrooke, and J.O. Von Uexkull, J. 1934. A stroll through the 1994b. Tolerance of bitter compounds by Schmidt. 1989. The detoxification of ant worlds of animals and men: a picture book an herbivore, Cavia porcellus. Journal of (Pogonomyrmex) venom by a blood fac- of Iinvisible worlds. Pp.5-76 (translation) Chemical Ecology 20:303-308. tor in horned (Phrynosoma). in C.H. Schiller, ed. 1957. Instinctive Be- Norman, D.M., J.R. Mason, and L. Clark. Copeia 3:603-607. havior: The Development of a Modem 1992. Capsaicin effects on consumption Semel, B. and L.K. Nicolaus. 1992. Estro- Concept. International Universities Press, of feed .by cedar waxwings, and house gen-based aversion to eggs among free- New York. finches. Wilson Bulletin 104549-551. ranging raccoons. Ecological Applications K.K. Wagner and M.R. Conover. 1999. Ef- Pelchat, M.L., H.J. Grill, P. Rozin, and J. 2:439-449. fect of preventive coyote hunting on sheep Jacobs. 1983. Quality of acquired re- Shivik, J.A., and D.J. Martin. In press. Aver- losses to coyote predation. Journal of sponses to tastes by Rattus norvegicus de- sive and disruptive stimulus applications Wildlife Management 63(2):606-6 12. pends on the type of associated discom- for managing predation. Proc. Eastern Werner, S., A. El Hani, and J.R. Mason. fort. Journal of Comparative Psychology Wildlife Damage Management Confer- 1998. Repellent coatings for irrigation 97: 140-153. ence. hose: effectiveness against coyotes. Jour- Reidinger, R.F. and J.R. Mason. 1983. Sterner, R.T. and S.A. Shumake. 1978. Bait- nal of Wildlife Research 2: 146-148. Evaluation and exploitation of weaknesses induced prey aversions in predators: some Wickler, W. 1968. Mimicry in Plants and in behavioral defenses against dietary poi- methodological issues. Behavioral Biol- Animals. New York, McGraw-Hill Pub- soning. Pp. 20-39 in D.E. Kaukienen, ed. ogy 22565-566. lishing Company, New York. Test Methods for Vertebrate Pest Control Stevens, G.R., J. Rogue, R. Weber, and L. Witmer, G.W., J.L. Bucknall, T.H. Fritts, and and Management Materials. ASTM STP Clark. 2000. Evaluation of a radar-acti- D.G. Moreno. 1996. Predator manage- 817, American Society for Testing and vated demand-performance bird hazing ment to protect endangered avian species. Materials, Philadelphia. system. International Biodeterioration and Transactions of the North American Wild- Riley, A.L., and D.L. Tuck. 1985. Conditioned Biodegradation 45: 129-137. life and Natural Resources Conference food aversions: a bibliography. Pp. 381-437 Swihart, R.K. 1990. Quebracho, thiram and 61: 102-108. in N.S. Braveman and P. Bronstein, eds. methiocarb reduce consumption of apple Witmer, G.W. and M.W. Fall. 1995. Man- ExperimentalAssessments and Clinical Ap- twigs by meadow voles. Wildlife Society aging mammalian carnivores to protect en- plications of Conditioned Food Aversions. Bulletin 18: 162-166. dangered species (abstract). The Wildlife Annals of the New York Academy of Sci- Ternent, M.A. and D.L. Garshelis. 1999. Society 2ndAnnual Conference, Portland, ences Series, Volume 443. NewYorkAcad- Taste-aversion conditioning to reduce nui- OR. emy of Sciences, New York. sance activity by black bears in a Minne- Zemlicka, D. and J.R. Mason. 2000. Re- Sacks, B.N., K.M. Blejwas, and M.M. Jae- sota military reservation. Wildlife Society sponse of captive coyotes to Renardine ger. 1999. Relative vulnerability of coy- Bulletin 27:720-728Ti11, J.A. and F.F. coyote repellent. Proceedings of the Ver- otes to removal methods on a northern Knowlton. 1983. Efficacy of denning in tebrate Pest Conference 19:336-338.

Vol. 18 No. 4 2001 Endangered Species UPDATE 181

Disadvantages of guarding ing the predator, herding the sheep, use, but sheep mortalities did not dif- dogs include: some dogs not stay- or positioning themselves between fer between producers with (n = 35) ing with or harassing sheep; some the sheep and predator. and without (n = 32) llamas during dogs, especially Komondors, being Franklin and Powell (1993) sur- the second year in Utah. overly aggressive toward people veyed 145 producers that used lla- (Green and Woodruff 1988; Andelt mas, primarily in Montana, Wyo- Donkeys 1992); and the dogs can be subject ming, Colorado, California, and Or- The National Agricultural Statistics to injury and premature death. egon. Most producers used one Service (2000) reported that 9% of Many of the disadvantages are rela- gelded male with 250 to 300 sheep sheep producers in the United States tively uncommon. Most producers in 250 to 300 acre pastures. Nearly used donkeys (Equus asinus) to pro- surveyed feel strongly that the ad- all llamas were not raised with tect sheep from predators during vantages of their dogs far outweigh sheep and were not trained to guard 1999. Donkeys apparently have an the disadvantages. sheep. One llama was more effec- inherent dislike for dogs and other Green and Woodruff (1988) re- tive than multiple llamas for deter- canids. They will bray, bear their ported that the rate of success in ring predation; the effectiveness of teeth, run and chase, and attempt to protecting livestock from predators gelded males, intact males, and fe- bite and kick an intruder (Green 1989). did not vary among Great Pyrenees, males was similar. Producers re- Donkeys apparently are most Komondor, Akbash, Anatolians, ported, however, more problems effective in small open pastures or Maremma, and hybrids, nor was the with intact (25% of 61 intact males) where sheep graze together. Green rate of success different among than gelded males (5% of 135 (1989) and Walton and Feild (1989) males and females or intact and gelded males) attempting to breed recommended using only one jenny neutered dogs. Dogs that were with ewes. Sheep that were intro- or gelded jack per pasture because reared with livestock from at most duced to llamas in corrals initially two or more donkeys often stay to- two months old, however, had a sig- sustained lower mortality than those gether instead of being with the nificantly higher rate of success introduced in pastures. The success sheep. Intact jacks generally are than dogs that were older than two of llamas was not related to age when too aggressive around sheep. Don- months when placed with livestock. the llama was introduced, age of keys generally should be allowed Andelt (1999) reported that ratings llama (after one or two years old) four to six weeks for bonding with of the effectiveness of guarding when guarding, if lambs were present sheep before they are used to deter dogs by producers using one breed or absent when the llama was intro- predators. Donkeys should be re- of dog in Colorado did not differ duced, or between open and covered moved during lambing because among breeds, but producers that (forested, shrub lands, gullies, ra- they might trample lambs or disrupt used multiple breeds rated Akbash vines, etc.) habitat. the ewe-lamb bond. Green (1990) more effective than Great Pyrenees Franklin and Powell (1993) re- recommended challenging a don- and Komondors. ported that gelded male llamas cost key with a dog to test its response $700 to $800, whereas intact males to canids; donkeys that are not ag- Llamas were about $100 less. Most produc- gressive should not be used. Llamas have been used to deter pre- ers reported that daily care for llamas Walton and Feild (1989) re- dation primarily by coyotes, red was the same as for sheep and that ported that the average purchase foxes (Vulpes vulpes), and dogs no special feeds were provided. Av- price per donkey was $144. They since the early 1980s. The National erage annual expense was $90 for also reported that average annual Agricultural Statistics Service feed (not including pasture) and vet- upkeep per donkey was $66. (2000) reported that 13% of the erinary costs were about $15. sheep producers in the United Franklin and Powell (1993) re- Bonding sheep and goats to States used llamas to protect sheep ported that 21% of ewes and lambs cattle from predators during 1999. Lla- were killed annually before acquir- Bonding young sheep to cattle mas are naturally aggressive toward ing a llama and 7% afterwards. (Anderson et al. 1987; Hulet et al. coyotes and dogs. Typical re- Meadows and Knowlton (2000) re- 1987) and goats to sheep that have sponses of llamas to coyotes and ported that producers with llamas lost been bonded to cattle (Hulet et al. dogs are being alert, alarm calling, significantly fewer sheep (n = 42) to 1989) has reduced coyote preda- walking to or running toward the predators than producers without lla- tion. This technique has not been predator, chasing, kicking, or paw- mas (n = 128) during the first year of readily adopted by sheep producers,

Vol. 18 No. 4 2001 Endangered Species UPDATE 183 possibly because of the additional mas are afraid of bears and moun- tridges for denning coyotes, and labor and expense involved with tain lions. Guarding dogs also ap- livestock protection collars have bonding sheep and goats to cattle. peared successful in protecting been restricted or eliminated by bal- cattle from wolf predation lot initiatives in some states such Relative effectiveness of (Coppinger et al. 1988), and were as Arizona, California, Colorado, guarding animals fairly effective in keeping wolves and Massachusetts (Manfred0 et al. Benefits of using guarding animals and black bears from carrion feed- 1999). The public also has rated include a decrease or elimination of ing sites in Minnesota (Coppinger guarding animals as more accept- predation, reduced labor to confine et al. 1987). able than most other techniques for sheep at night, more efficient use Several methods, including reducing predation (Arthur 1981; of pastures for grazing, reduced re- livestock confinement, disposal of Reiter et al. 1999). Thus, guarding liance on other predator control livestock carcasses, herders, fenc- animals are one of the few remain- techniques, and a greater peace of ing, frightening devices, trapping, ing successful techniques, in some mind. A comparison of surveys snaring, sodium cyanide ejectors, states, that livestock producers can where producers reported the aver- "denning" (locating the dens of use to mitigate predation. age annual value of sheep saved per depredating coyotes and killing the Green and Woodruff (1989) and guarding animal suggests guarding pups andlor adults), aerial hunting, Green et al. (1993) reported that dogs, compared to llamas, saved ground shooting, hunting with de- guarding dogs repelled black bears more sheep from predators (Table coy dogs, livestock protection col- and grizzly bears (Ursus arctos) 1). Guarding dogs and llamas have lars, and poison baits have been during most encounters, and Andelt been rated as more effective than used to reduce predation on live- and Hopper (2000) reported that donkeys for deterring predation. stock (Andelt 1996). Poison baits producers with guarding dogs, The above comparisons should be were withdrawn from use in 1972 compared to producers without interpreted conservatively because (Andelt 1996), and use of some guarding dogs, sustained fewer ewe guarding dogs, llamas, and donkeys methods such as trapping, snaring, and lamb mortalities to black bears. were not surveyed in the same stud- sodium cyanide ejectors, gas car- Thus, guarding dogs may offer ies nor under the same sheep man- agement conditions. Advantages of donkeys and lla- Table 1. Average annual value of sheep saved from predation by mas over guarding dogs include: each livestock guarding animal and ratings of effectiveness of guard- less prone to accidental death; ing animals as reported in various studies. longer-lived; do not require special feeds; stay in the same pasture as sheep; apparently do not need to be raised with sheep; more compatible with other depredation control tech- niques such as traps, snares, M-44s, and livestock protection collars; and donkeys are cheaper than guarding dogs. Guarding dogs ap- parently have some advantages over donkeys and llamas. Guard- ing dogs deter predators in fenced pastures and on open range, whereas llamas and donkeys appear most effective in fenced pastures smaller than 300 acres. Guarding "reen et al. (1 984) dogs are effective in deterring bear bFranklin and Powell (1993) and mountain lion predation (Green "ndelt and Hopper (2000) dGreenand Woodruff (1 988) and Woodruff 1989; Andelt and eAndelt (1 992) Hopper 2000), whereas some don- 'Meadows and Knowlton (2000) keys (Green 1989) and possibly lla- gWalton and Feild (1989)

184 Endangered Specks UPDATE Vol. 18 No. 4 2001 some potential for reducing grizzly Andelt, W.F. 1999. Relative effectiveness Green, J.S., R.A. Woodruff, and T.T. bear predation on livestock, which of guarding-dog breeds to deter preda- Tueller. 1984. Livestock-guarding dogs tion on domestic sheep in Colorado. for predator control: costs, benefits, and may result in conserving more bears. Wildlife Society Bulletin 27:706-714. practicality. Wildlife Society Bulletin Many of our carnivores are Andelt, W.F., and S.N. Hopper. 2000. 12:44-50. found on private lands and in areas Livestock guard dogs reduce predation Green, J.S., R.A. Woodruff, and P.J. Wick. where livestock are grazed on pri- on domestic sheep in Colorado. Journal 1993. Bears, ostriches, and specialized vate and public lands. Producers of Range Management 53:259-267. grazing: putting guarding dogs to work. Anderson, D.M., C.V. Hulet, J.N. Smith, Proceedings of the Great Plains Wildlife need to have successful techniques W.L. Shupe, and L.W. Murray. 1987. Damage Control Workshop 11: 105-108. for resolving predation on live- Bonding young sheep to heifers. Ap- Hulet, C.V., D.M. Anderson, J.N. Smith, stock. Without successful methods, plied Animal Behaviour Science 19:31- and W.L. Shupe. 1987. Bonding of predation on livestock will increase 40. sheep to cattle as an effective technique Arthur, L.M. 1981. Coyote control: the for predation control. Applied Animal and animosity by livestock produc- public response. Journal of Range Man- Behaviour Science 19: 19-25. ers toward predators likely will also agement 34: 14-15. Hulet, C.V., D.M. Anderson, J.N. Smith, increase, which will result in fewer Coppinger, R., L. Coppinger, G. Langeloh, W.L. Shupe, C.A. Taylor, Jr., and L.W. attempts to conserve carnivores. and L. Gettler. 1988. A decade of use Murray. 1989. Bonding of goats to Without successful techniques, pro- of livestock guarding dogs. Proceedings sheep and cattle for protection from Vertebrate Pest Conference 13:209-214. predators. Applied Animal Behaviour ducers also may sell their land for Coppinger, R., J. Lorenz, and L. Science 22:261-267. other uses, such as subdivisions, Coppinger. 1983. Introducing livestock Manfredo, M.J., C.L. Pierce, D. Fulton, J. which are less conducive to carni- guarding dogs to sheep and goat produc- Pate, and B.R. Gill. 1999. Public ac- vore conservation. Thus, guarding ers. Proceedings Eastern Wildlife Dam- ceptance of wildlife trapping in Colo- age Control Conference 1:129- 132. rado. Wildlife Society Bulletin dogs, llamas, and donkeys should Coppinger, R., J. Lorenz, and L. 27:499-508. be thought of as valuable additions Coppinger. 1987. New uses of livestock Meadows L.E., and F.L. Knowlton. 2000. to the toolbox of management prac- guarding dogs to reduce agriculture1 Efficacy of guard llamas to reduce ca- tices that reduce predation on live- wildlife conflicts. Proceedings Eastern nine predation on domestic sheep. Wild- stock. In addition, use of guarding Wildlife Damage Control Conference life Society Bulletin 28:614-622. 3:253-259. National Agricultural Statistics Service. animals may result in enhanced Franklin, W.L., and K.J. Powell. 1993. 2000. Sheep and goats predator loss. conservation of carnivores. Guard llamas. State University Ex- USDA, National Agricultural Statistics tension Bulletin, Ames, Iowa. Service, Report Mt An 2 (5-OO), Wash- Literature cited Green, J.S. 1989. Donkeys for predation ington, D.C. Andelt, W.F. 1985. Livestock guarding control. Proceedings Eastern Wildlife Pearson, E.W. 1986. A literature review dogs protect domestic sheep from coy- Damage Control Conference 4:83-86. of livestock losses to predators in west- ote predation in Kansas. Proceedings Green, J.S. 1990. Reducing predation with ern U.S. Final Report, U.S. Department Great Plains Wildlife Damage Control guarding animals. University of Cali- of Interior, Fish and Wildlife Service, Workshop7:lll-113. fornia Hopland Field Station Publication Denver, Colorado. Andelt, W.F. 1992. Effectiveness of live- 101:62-68. Pfeifer, W.K., and M.W. Goos. 1982. stock guarding dogs for reducing preda- Green, J.S., and R.A. Woodruff. 1988. Guard dogs and gas exploders as coyote tion on domestic sheep. Wildlife Soci- Breed comparisons and characteristics of depredation control tools in North Da- ety Bulletin 20:55-62. use of livestock guarding dogs. Journal kota. Proceedings Vertebrate Pest Con- Andelt, W.F. 1995. Livestock guard dogs, of Range Management 41:249-251. ference 10:55-61. llamas, and donkeys for reducing live- Green, J.S. and R.A. Woodruff. 1989. Live- Reiter, D.K., M.W. Brunson, and R.H. stock losses to predators. Cooperative stock-guarding dogs reduce depredation Schmidt. 1999. Public attitudes toward Extension Bulletin Number 1.218, Colo- by bears. Pp 49-54 in M. Bromley, ed. wildlife damage management and policy. rado State University, Fort Collins. Bear-people conflicts: proceedings of a Wildlife Society Bulletin 27:746-758. Andelt, W.F. 1996. Carnivores. Pp133- symposium on management strategies. Walton, M.T. and C.A. Feild. 1989. Use 155 in P. R. Krausman, ed. Rangeland Northwest Territories Department of Re- of donkeys to guard sheep and goats in wildlife. The Society for Range Man- newable Resources, Yellowknife, North- Texas. Proceedings Eastern Wildlife agement, Denver, Colorado. west Territories, Canada. Damage Control Conference 4:87-94.

Vol. 18 No. 4 2001 Endangered Species UPDATE 185 Opinions Historical Attitudes and Images and the Implications on Carnivore Survival Karlyn Atkinson Berg Wolf Consultant, Humane Society of the United States, 44781 Bittner Point Road, Bovey, MN 55709; (218) 245-3049; [email protected]

Abstract This paper examines how mythological images and historical attitudes emerge and influence our interactions with different predator species, such as the grizzly bear (Ursus arctos), cougar (Puma concolor), lynx (Lynx canadensis), wolf(Canis lupus), coyote (Canis latrans), and raven (Corvus corax). The author will compare the relationship between humans and carnivores, and how attitudes and beliefs have impacted different predator species. Do people regard certain carnivores as more fierce, dangerous, or problematic? Is there more animosity and disparate levels of hostility or tolerance toward the different carnivores? Have these attitudes influenced concepts and ethics applied to wildlife management? How is the value of predators measured, considered or applied? Can understanding the different perceptions help resolve complicated issues, such as reintroduction, critical habitat, depre- dation conflicts, animal damage control, and management? The author believes scientific knowledge is not enough to achieve acceptance of carnivores. The purpose of this inquiry will be to discover if knowledge and education can develop understanding and tolerance of all predators, and thus enhance the commitment to co-exist with carnivore species

Mythlcal images and historical attitudes Historical attitudes and cultural threat to humans, but had more to may still influence human interactions beliefs have impacted many differ- do with deeply rooted bias and with carnivores such as the grizzly bear ent predator species. From Viking mythological symbolism. (Ursus arctos), cougar (Puma bearskin wearers to the arrival of The cougar is commonly de- concolor), lynx (Lynx canadensis), humans in the New World, most scribed as the coward. Theodore wolf (Canis lupus), coyote (Canis carnivores continued to meet the Roosevelt described a cougar he latrans), and raven (Cowus corax). same fate. Most of the legends re- had treed as "the big horse-killing The earliest historic records, creation volve around the fear of wilderness, cat, the destroyer of the deer, the stories, and fables were examined. No- the idea of good and bad animals, and lord of stealthy murder, facing his tably, at the advent of agriculture, the need to remove all that stands in doom with a heart both craven and Akkadian literature delineates the split the way of progress. Species with cruel" (Danz 1999; Worster 1977; between humans and nature, and the fang and claw that hunted the good Roosevelt 1913). The cougar is re- god Ea predicts that from this time for- prey (which humans wanted for peatedly described as a cunning, ward, nature will be hostile to man. He themselves) are traditionally de- merciless, and sneaky cat. The por- asserts that lion, wolf, famine and scribed as the bad animals. The trayal is made with little concern plague will not be removed from predator is reduced to the status of that a trait like stealth is a neces- humankind's dilemmas, but provided marauder and thief, and hence sub- sary ability to survive as a cougar, "to rise up and cut the people low" jected to extermination. and has little to do with intentions (Gardnerl984). This division between Do people regard certain carni- of mere cruelty for cruelty's sake. wilderness and the tamed domestic vores as more fierce, dangerous, or Predation is often perceived as lands that humans seek does seem to problematic? Some species have murder, not as a pursuit of food. have taken place, and remains a cur- been described with more vivid The bear, however, is character- rent conflict among predators, wilder- hostile imagery that did not reflect ized more in terms of admiration, ness advocates, and ranchers. actual biological evidence of their combining descriptions of its sav-

186 Endangered Species UPDATE Vol. 18 No. 4 2001 ing, "A grizz or a brown is a bear, but a black is a dog. They don't even belong in the same category. I don't have much respect for a black bear as a vicious animal" (Kaniut 1983). Native Americans admired the per- severance and hunting prowess of the wolf. These traits were despised by wolf hunters who consistently describe the wolf as ruthless, treacherous, and a cruel demon that exists in spite of man's will. Myths also accuses the wolf of not only devouring human bodies, but also devouring the soul for the devil. The "yellow eyes as the lamps of the devil" is apparently still an image that persists (Berg un- published manuscript). A Minnesota state legislator recently exclaimed, "wolves are serial killers" (McAuliffe 1999), and a sign held at hearings at declared, "Pedophilesand wolves kill children" (Sandstrom 1998) The coyote is described as a te- nacious varmint (the last of the out- laws), detested for its intelligence, and defined as crafty or the mytho- logical Trickster. As prophesied for the wolf, fears are exaggerated about predators, and people pro- claim that it is only a matter of time Grizzly bear (Ursus arctos). Photograph by B. Moose Peterson1 before a coyote is going to jump WRP. onto someone's deck, grab their child, and run off. (Interestingly, a proposed black bear hunt in New Jersey was canceled after wildlife organizations protested. But there was minimal outcry over the coy- ageness with the animal's almost same trails, fished for the same ote hunt that was also proposed.) human dignity. Descriptions such salmon, dug for roots, harvested People see the bear as a cuddly ani- as "unbelievable size of the brute" berries, seeds, and nuts, caught mal worth protecting because of and "lordly intelligence" (Young meat, and could stand upright like their teddy bear experience. Con- 1980) serve as examples of the humans. While the grizzly bear was versely people fear the wolf and bear's more dignified status among feared as dangerous, it was consid- coyote because of the negative the carnivores. The name Old ered a powerful shaman of the ani- childhood stories about these ca- Ephraim, taken from the Biblical mal world. The depiction of the nines. Unfortunately the Teddy patriarch of noble lineage, is hon- bear is not infamous, though it was Bear image causes problems be- orably bestowed upon the bear. considered dangerous for humans cause people feel they can treat real Other cultures, such as the Na- to encounter. Keith Johnson, a mas- black bears as the cute stuffed bear tive Americans, viewed bears as ter guide, actually admires the al- they have at home. This ultimately half human, for the bear walked the leged viciousness of the bear, say- puts the people and bears in danger.

Vol, 18 No. 4 2001 Endangered Specles UPDATE 187 Mythical depiction has not predators was not an accidental per wolf of 1438 wolves living pri- spared any of these species. Even byproduct of progress, but a well- marily in the wilderness, and an ad- the lordly bears or monarchy of the organized extermination. The BBS ditional $197 per wolf outside the big cats were considered killers in was also a center for state bounty designated area (surplus animals, the way of progress. The value of information, which published pam- dismissed as unnecessary extras). a varmint has traditionally been cal- phlets on the habits of predators, Unfortunately, this study culated as being worthless. The and bulletins about the best trap- wrongly assumed the cost of the historic, negative attitudes prevail; ping scents and poisons with which compensation program were pay- throughout history, humans have to kill the animals. In 1915, Con- ments made to verified wolf depre- not progressed, learned from mis- gress appropriated $125,000 di- dation cases only. In fact, the com- takes, improved methods, nor be- rectly to fund this war on predators. pensation granted by state conser- come enlightened on the value of This amount kept increasing; thus vation officers is not based on sci- predators. Unfortunately, predator in 1950, $1,098,00 was allocated, entific verification, and often con- programs and proposals of the year and by 1970, $3,267,000 (Advisory flicts with the Federal examination 2000 are not much improved from Committee on Predator Control findings. Considerable payments those of the past. Society seems to 1972). The program was defined were made to cases that were not be in an era of rational compromise by Jenks Cameron of the BBS as wolf-related or verified, and some that may be headed in reverse. "suppressive warfare against unde- were documented by the Federal Perhaps it has been forgotten sirable, injurious wildlife," and "the program as outright fraud. that conservation evolved from the protection and encouragement of The thesis also wrangly as- progressive political movement wildlife in its desirable and benefi- sumes that most or all wolves out- which valued nature, lands, trees, cial forms" (Worster 1977). side of the designated area contrib- and wildlife as a commodity to be In these modern times, most of ute to depredation. Current wolf used for human economic success. us have thought that the idea of poi- plans appear to be determined by The movement set forth the goals son, especially 1080, would not how many wolves humans can kill of more efficient management and loom again over wildlife, or that and get away with, rather then how maximum utilization of natural re- bounties-repeatedly proven to be few wolves create problems and sources, not conservation or the ineffective, fraudulent, and wasteful need to be removed. The study preservation of biodiversity. Be- -would not be promoted. However, wrongly excludes the savings that tween 1901 and 1909, as the pro- Wyoming recently proposed a $1000 could occur with good, preventative gressive ideology developed, it bounty, fortunately vetoed by the husbandry methods, set in place soon included an extensive program governor. But in Minnesota, Gov- with non-lethal advancements. designed to exterminate predators ernor Jesse Ventura just signed a However, is the only measure of and make America safe from their law that includes a $150 bounty, their existence based on a bargain thieving presence. The rescue of now called a predator payment. The price value system? E.O. Wilson says nature from these killers would no new management law also allows dis- it is prudent "tojudge every scrap of longer be carried out just by pio- cretionary killing by ranchers. biodiversity as priceless" (Wilson neers, ranchers, and lone wolf hunt- People are familiar with legis- 1992). What is the real cost if wolves ers. The new Bureau of the Bio- lators' negative views on predators, are not allowed to exist? logical Survey (BBS), as part of the but what about the modern scien- It is most distressing that a bi- Department of Agriculture, would tists? Have their data redefined the ologist should calculate the eco- now reinforce the ferocity against value of the varmint by its ecologi- nomic value of wolves, rather than predators. Thus, a major focus of cal place in the ecosystem? It is focusing on their ecological value, conservation in the early years was quite disappointing to see recent by saying that "wolves inhabiting the well-funded, staunch effort to scientific papers calculating mon- wilderness cost little to societyt' deliberately eradicate predators. etary formulae to promote the view (Wilson 1992). Wilson's words Today, this practical technique of that it is cost efficient to only main- seem to fit this sad situation: "If a killing any creature that does not tain a population of 1400 wolves in price can be put on something, that suit humans has been renamed flex- the wilderness and semi-wilderness something can be devalued, sold ible management. of Minnesota (e.g., Mech 1998). and discarded" (Wilson 1992). It is important to remember that The average annual cost, according The study ponders an even the tremendous loss of our nation's to these calculations, would be $86 more disturbing premise by noting

188 Endangered Species UPDATE Vol. 18 No. 4 2001 that trapping and hunting would not wilderness, or Disney areas with The future of carnivores is still threat- be efficient methods of wolf con- just enough species to entertain ened by hostile attitudes. Wild car- trol. The following question is from a safe distance? Is a wilder- nivores are a part of American cul- raised: Without using poison or ness experience a mob howling ad- ture and heritage; their preservation substantial financial incentives, venture? Do carnivores need to fit cannot be neglected. Responsible would the Minnesota wolf population into approved behaviors? Are stewards need to be on a path of co- be controllable in 2005? It is pro- people willing to not intrude some existence, not returning to the sanc- posed that the sooner controls begin, habitats, or at least enter with tioned destruction of America's the easier and less costly it will be. knowledge and respect? Do carni- predators. Sadly, society seems to have vores have an intrinsic right to exist entered a monumental era derived without directly helping humanity? Literature cited from the philosophy of Descartes. The majority of Americans sup- Advisory Committee on Predator Control. Apparently, all nonhuman animals port protection of large predators 1972. 1971 Predator control report to the Council on Environmental Quality and the are machines, and nature (as a gi- and restoration of a diverse, natu- Department of the Interior. Institute for ant mathematical mechanism) is to ral ecosystem. They do not support Environmental Quality and United States be measured, counted, computer- the abuse and over-utilization of Department of the Interior University of ized, and placed on a GIs map, with public lands, nor the destruction of Michigan, Ann Arbor, Michigan. a statistical and monetary formula wild animals, wild lands, and natu- Berg, K. 1983. Wolves and Humans His- tory. Unpublished manuscript. to determine how many specimens ral resources for private interest Berg, C. 1999 Minnesota Senate wolf are profitable and thus allowed to groups. Humans have a responsi- Hearings, St Paul. Minnesota survive. It seems that humans have bility to protect other species, even Danz, H.P. 1999. Cougar. Swallow Press/ yet to find what Aldo Leopold if it is inconvenient. Species should Ohio University Press, Athens, Ohio. Gardner, J. and J. Maier. 1984. Gilgamesh. called ecological conscience not be removed from protection for Alfred Knopf, New York. (Leopold 1949). As long as preda- short term political favors to spe- Kaniut, L. 1983. Alaska BEAR Tales. tor control is based on the bullet as cial interest groups. The stakehold- Alaska Northwest Books, Anchorage, the cheapest method, there will be ers of our lands have been narrowly Alaska. no earnest attempt to initiate pre- defined as consumptive users for Leopold, A. 1949. A Sand County Almanac. Oxford University Press, New York. ventative, non-lethal, and humane too long. More importantly, future McAuliffe, B. 1999. Wolf bill may be management programs. generations and wild creatures are dead. Minneapolis Star Tribune, April Fear is a force still working the most vulnerable to today's ac- 15. Minneapolis, MN. against predators. Although some tions. What legacy will society Mech, L.D. 1998. Estimatedcosts of main- taining a recovered wolf population in carnivores are capable of attacking leave for future generations? agricultural regions of Minnesota. Wild- people, these are rare encounters. Carnivore recovery should be life Society Bulletin 26(4):817-822. Education has to include not only based on ecological principles, not Roosevelt, T. 1913. A cougar hunt on the how to avoid predator encounters, political pandering. Recovery must rim of the Grand Canyon. Outlook 105: but also teaching a willingness to ensure the long-term survival of the 259-266. Wilson, E.O. 1992. Diversity of Life. give them space. It has been clearly species, and also restore functional BelknaptHarvard University Press, demonstrated that modern tech- ecosystems that will benefit the full Cambridge, Massachusetts. nologies that decimate nature are range of wildlife and human com- Worster, D. 1977. Nature's Economy. Si- adequate. Society needs to strive munities that depend on them. erra Club Books, San Francisco. Young, EM. 1980. Encounters of Lewis and and find ways for living success- Coexistence with large predators Clark to Modern Times: Man Meets Griz- fully in association with our native requires tolerance and education. zly. Houghton Mifflin Publishers, Boston. fauna and flora. Do people want

Vol. 18 No. 4 2001 Endangered Species UPDATE 189 Bringing Down the Walls

Gale H. Ford Executive Director, Grizzly Discovery Center; West Yellowstone, MT; (406) 646-7001 ; (fax) (406) MTBEARDOCQaol.com

Educational venues to view, interact, cation of young people. Alternative same experience as hearing a pack of and learn about wildlife occur in avenues for inspiring an interest in wolves in Lamar Valley, the zoo still many settings, including zoos, pre- wildlife and wild lands must be rec- allows the visitor to have a close and serves, refuges, rehabilitation centers, ognized and enhanced for their abil- personal moment with this species. national parks, and other wild lands. ity to establish a closer relationship The next headline about a wolf is Each human believes his or her venue between people and wildlife. more likely to be read by someone or political camp to be the best model The need for effective conserva- who knows the sound of a howl than for teaching ecology. The world is tion education is absolutely urgent to- one who does not. Even more im- rapidly changing, both in the effects day. The desire to understand or portantly, the impact on wild habi- created by increasing human popu- merely view animals has been joined tats should be considered if human- lations, and the often-heroic attempts by the need to conserve their popu- animal connections in a wild setting to halt and even reverse such trends. lations and ecosystems. Over the last were the only experiences allowed. In the 1960s, "ecology" was a new, few decades, the explosive growth of Openness to learning about new mysterious word to most people. human populations has been linked exhibition techniques, husbandry Thirty years later, the term "conser- with changes in the abundance of programs and educational outreach vation" has been expanded beyond a other species. The understanding of programs offered in today's zoologi- resource management definition and animals' needs and of human impacts cal institutions needs to be embraced now includes concern with retaining on animal populations became criti- as a viable non-consumptive aug- the planet's biodiversity and the ac- cal; the future of these animals de- mentation to wildlife education. For tions that affect such change. pends on people's actions. This un- the general visitor, zoos and aquari- These concerns for sustainability, derstanding is no longer optional. ums serve the conservation commu- cessation of extinction, and enhance- Everyone in this discipline knows the nity by providing information in a ment of the welfare of endangered message, but in spite of its size, this relaxed, family-oriented setting. species ultimately will succeed only group is still in the minority. Zoos and aquariums are positioned if they do not fall on deaf ears, but Today, conservation education to play an ever-changing role in con- on the ears of those who know the needs to be recognized as a specialty servation through education. language and arena of concern. As that benefits from diverse technolo- Conservation requires an ex- the family structure has taken on a gies, varied educational efforts, and panding perspective, but educators different face from that of the past, multi-disciplinary expertise. The are divided and confounded by dis- the connection between young people conservation message needs to shift agreements about the message and and wildlife must be nurtured and not from merely endangerment, captive the appropriate media of delivery. left to chance. Parents and grandpar- breeding, or even reintroduction to Conservationists must share united ents often were the inspiration for include and emphasize the impor- intentions in order to be effective. wildlife wonderment through shared tance of saving habitat. The exper- Advocacy groups, wildlife protection outings into the wild. Each person tise to facilitate the delivery of this groups, and zoological educators can probably recall those mentors message exists. It is present within share a common goal: to instill in oth- who were the frst to introduce nature. conservation groups and in areas that ers the passion to protect, conserve, Today's living and working con- were previously ignored. restore, and ultimately live harmoni- ditions do not foster such opportuni- Zoos and aquariums draw 121 ously with the unique wildlife and ties. Urban living situations, in- million visitors each year. Captive wild lands in the environment. creased job demands, widening eth- wildlife facilities have historically Therefore, the crucial partnerships nic perspectives, an aging population, been considered substandard or poor that make the conservation message and changes in nuclear family struc- substitutes for wildlife experiences. reach the most ears, in the most ef- ture are but a few of the major shifts While hearing wolves howl at the fective manner humanly and hu- drastically affecting the nature edu- Grizzly Discovery Center is not the manely possible, should be fostered.

190 Endangered Species UPDATE Vol. 18 No. 4 2001 News from Zoos

Minnesota Zoo, China Work to Save World's Most Endangered Tiger

Minnesota Zoo Conservation Director Dr. Ron Tilson led a team of Chinese conservation officials on an expedition through Longyan, Fujian Province, in an effort to save the critically endangered South China Tiger. Dr. Tilson also held a tiger conservation workshop, which marks the beginning of a partnership between China and foreign officials, including AZA's Tiger Species Survival Plan, on the South China Tiger Project. More than 20 conservation officers from seven provinces attended the workshop, which highlighted tiger track- ing and search techniques. Following the expedition, the team visited the South China Tiger Captive Breeding Program in Meihuashan Mountain, where "first phase facilities" have been constructed to breed and prepare the tigers for release. The South China Tiger, know to locals as a "Mountain God," is the most endangered subspecies of tigers know to exist. Fewer than 30 live in the wild and only 60 are held in Chinese Zoos. [Source: Communique]

Namibian Cheetah Ambassadors Come to the States

Ten young Namibian cheetahs arrived in the United States in April for distribution to AZA accredited zoos who participate in the Cheetah Species Survival Plan (SSP). These cheetahs, comprising a group of cats from eight different litters, all originated from commercial farmland areas and eventually found sanctuary at the Cheetah Con- servation Fund (CCF). The cheetahs represent a presidential gift from Namibia to the United States in recognition of the support it has given to cheetah conservation efforts in Namibia. Four of the cheetahs are housed at the Cincinnati Zoological Gardens and the other six were given to the White Oak Conservation Center. They will be integrated into the Cheetah SSP, which was established in 1982 and manages of all captive cheetahs at different facilities in North America as a unit. Working within this program, all the facilities holding cheetahs cooperate on issues concerning reproduction, genetics, diets and general husbandry of the species. CCF selected these ten cheetah based strictly on the criteria that they are non-releasable animals. The circum- stances under which these cheetahs were orphaned are mostly tragic, with their mothers being shot. However, all of the cubs found their way to CCF through the concern for their welfare of the people that caught them and in two cases, through the speedy intervention of a concerned neighbor. CCF has been the recipient of three grants from the AZA's Conservation Endowment Fund. [Source: Jack Grisham, AZA Cheetah SSP Coordinator]

Mexican Gray Wolves Make Zoo New Home

The National Zoo announced that it has made a home for three Mexican gray wolves as part of an international recovery effort to reintroduce the highly endangered wolf subspecies into the wild in Arizona, New Mexico and Mexico. At a news conference this morning at the new wolf exhibit, located uphill from the seal pool, zoo and animal conservation officials said the wolves will stay at the zoo until they can be released into a wilderness area in either Arizona, New Mexico or Mexico. The National Zoo participates in the Mexican Wolf Species Survival Plan (SSP), which manages the Mexican wolves held in captivity. The SSP and the US Fish and Wildlife Service work cooperatively to support the Mexican Wolf Recovery Program, the international effort to reintroduce Mexican Wolves. The Mexican gray wolf is the most rare and genetically distinct subspecies of gray wolf in North America but have not been seen in the United States and Mexico since 1970 and 1980, respectively. There are about 200 Mexican gray wolves living today, and nearly all of them were born and raised in captivity. Conservation officials said they work with the ranchers to try to keep the wolves away from humans and live- stock, and they attach radio collars to the animals to track their whereabouts. [Adapted from an article by Karlyn Barker, The Washington Post]

Information for News from Zoos is provided by the American Zoo and Aquarium Association

Endangered Species UPDATE 191 Vol. 18 No. 4 2001