Journal of Mammalogy

David M. Leslie, Jr., Journal Editor, United States Geological Survey, Biological Resources Division, Oklahoma Cooperative Fish and Wildlife Research Unit, 404 Life Sciences West, Oklahoma State University, Stillwater, OK 74078-3051 (Direct all manuscripts to the Journal Editor); 405-744-6342; FAX 405-744-5006; [email protected] Ronald E. Barry, Associate Editor, Department of Biology, Frostburg State University, Frostburg, MD 21532- 1099; 301-687-4167; FAX 301-687-3034; [email protected] Troy L. Best, Associate Editor, Department of Biological Sciences, 331 Funchess Hall, Auburn University, AL 36849-5414; 334-844-9260; FAX 334-844-9234; besttro @ mail.auburn.edu Joseph A. Cook, Associate Editor, University of Alaska Museum, 907 Yukon Drive, Fairbanks, AK 99775- 1200; 907-474-6946; FAX 907-474-5469; [email protected] Meredith J. Hamilton, Associate Editor, Department of Zoology, Oklahoma State University, Stillwater, OK 74078-0459; 405-744-9685; FAX 405-744-7824; [email protected] John A. Litvaitis, Associate Editor, Department of Natural Resources, James Hall, University of New Hamp- shire, Durham, NH 03824; 603-862-2094; FAX 603-862-4976; [email protected] Brett Riddle, Associate Editor, Department of Biological Sciences, University of Nevada, Box 454004, Las Vegas, NV 89154-4004; 702-895-3133; FAX 702-895-3956; [email protected] Thomas E. Tomasi, Associate Editor, Department of Biology, Southwest Missouri State University, Spring- field, MO 65804-0095; 417-836-5169; FAX 417-836-6934; [email protected] C. Renn Tumlison, Associate Editor, Department of Biology, Henderson State University, Arkadelphia, AR 71999-0001; 870-246-5511; FAX 870-230-5144; tumlison @ hsu.edu Michael R. Willig, Editor for Special Features, Department of Biological Sciences, Texas Tech University, Lub- bock, TX 79409-3131; 806-742-2590; FAX 806-742-2963; [email protected] Kenneth T. Wilkins, Editor for Reviews, Department of Biology, Baylor University, Waco, TX 76798-7388; 254- 710-2911; FAX 254-710-2969; [email protected] Robert M. Timm, Productions Editor, Museum of Natural History, University of Kansas, Lawrence, KS 66045- 2454; 785-864-4180; FAX 785-864-5335; btimm @falcon.cc.ukans.edu Jill Franklin, Managing Editor, Allen Press Inc., 810 East 10th Street, Lawrence, KS 66044; 800-627-0629, ext. 284; FAX 785-843-1274; [email protected] Rod Lawrenz, Advertising Manager, Allen Press, Inc., 810 East 10th Street, Lawrence, KS 66044; 800-627- 0629, ext. 267; FAX 785-843-1274; [email protected]

Mammalian Speciesand Special Publications

Virginia Hayssen, Managing Editor for Mammalian Species and Special Publications, Department of Biological Sciences, Smith College, Northampton, MA 01063; 413-585-3856; FAX 413-585-3786; vhayssen @ science.smith.edu Elaine Anderson, cAssociate Editor for Mammalian Species, Denver Museum of Natural History, 2001 Colorado Boulevard, Denver, CO 80205-5798; 303-370-6470; FAX 303-331-6492 Leslie Carraway, Associate Editor for Mammalian Species, Nash 104, Department of Fisheries and Wildlife, Oregon State University, Corvallis, OR 97331; 541-737-1973; FAX 541-737-1980; [email protected] Serge Lariviere, Associate Editor for Mammalian Species, Ducks Unlimited Inc., Institute for Wetland and Waterfowl Research, One Waterfowl Way, Memphis, TN 38120-2351; 901-758-3825; FAX 901-758-3850; slariviere @ ducks.org Meredith J. Hamilton, Editor for Special Publications, Department of Zoology, Oklahoma State University, Stillwater, OK 74078-0459; 405-744-9685; FAX 405-744-7824; [email protected]

The JOURNAL OF MAMMALOGY (ISSN 0022-2372) is an official publication of the American Society of Mammalo- gists and is published quarterly in February, May, August, and November. Publishing services are provided by Alliance Communications Group, 810 East lOth Street, Lawrence, KS 66044. Calendar year subscription prices are: $35.00 for members and $170.00 for subscribers. Periodicals postage paid at Lawrence, KS. POSTMASTER: Send address changes to JOURNAL OF MAMMALOGY, Allen Marketing & Management, 810 E. lOth Street, P.O. Box 1897, Lawrence, KS 66044-8897; 785-843-1235; FAX 785-843-1274. ASM home page: www.mammalsociety.org and www.mammalogy.org @ 2000 The American Society of Mammalogists THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. Journal of Mammalogy, 81(2):462-47:

SELECTION OF MULE DEER BY MOUNTAIN LIONS AND COYOTES: EFFECTS OF HUNTING STYLE, BODY SIZE, AND REPRODUCTIVE STATUS

BECKY M. PIERCE,* VERNON C. BLEICH, AND R. TERRY BOWyER

Institute of Arctic Biology and Department of Biology and Wildlife, University of Alaska Fairbanks, Fairbanks, AK 99775 (BMP, VCB, RTB) California Department of Fish and Game, 407 West Line Street, Bishop, CA 93514 (BMP, VCB)

Predation on mule deer (Odocoileus hemionus) by mountain lions (Puma concolor) and coyotes (Canis latrans) was examined to test effects of hunting style and body size, and for mountain lions reproductive status, on selection of prey. Mountain lions, which hunt by stalking, selected :5l-year-old mule deer as prey. Body condition of mule deer did not affect prey selection by coyotes or mountain lions, and both predators preyed upon females and older adult deer more often than expected based on the percentage of these groups in the population. Female mountain lions selected female deer, but male mountain lions did not. Female mountain lions without offspring, however, did not differ from male mountain lions in prey selection. Coyotes did not select for young deer. Female mountain lions with kittens were selective for young deer in late summer.

Key words: California, Canis latrans, coyote, mountain lion, mule deer, Odocoileus hemionus predation, prey, Puma concolor

Differences in age, sex, and physical their ability to escape predation (Huggard condition may predispose parts of an un- 1993; Peterson 1977), and predators may gulate population to predation and cause kill animals in poor condition preferentially important changes in the demography and (Ackerman et al. 1984; Mech 1970) be- dynamics of prey (Curio 1976; Taylor cause selection for more vulnerable prey re- 1984). For example, selection by wolves quires less energy and poses less risk to the (Canis lupus) for older age classes of predator. Further, antipredator strategies of moose (Alces alces) on Isle Royale led to a prey may vary with group size, age, sex, younger population of moose but also con- and habitat use by prey (Bleich 1999; Bow- tributed to population fluctuations of both yer 1987; Karanth and Sunquist 1995). species (Mclaren and Peterson 1994). Sus- Large mammalian carnivores exhibit dif- ceptibility to predation may vary with size ferent styles of hunting. Some species use of predator (Bekoff et al. 1984; Huggard coursing tactics; others use stealth to stalk 1993; Ross and Jalkotzy 1996) or method and ambush prey (Kleiman and Eisenberg of hunting (stalking versus coursing- 1973). Coursing predators, such as wolves, Kruuk 1972; Kunkel et al. 1999). Ungulate may pursue moose for long distances populations are subject to predation by both (Mech 1970), apparently assessing moose canids and felids, and these predators often condition and the likelihood of a successful vary in body size and style of hunting kill (Peterson 1977). Wild dogs (Lycaon (Mech 1970; Packer et al. 1990; Schaller pictus) also pursue prey over great distanc- 1972). Physical condition of prey can affect es (Estes and Goddard 1967), and Kruuk * Correspondent:lionlady@ gateway.net (1972) noted that spotted hyenas (Crocuta

462 May 200C PIER( ETA! -PREDAnON BY MOUNTAIN LIONS AND COYOTES 463 crocuta) were more successful at killing 1981; Bowyer 1987), thereby increasing prey if the chase was > 300 m. Coyotes size of prey they kill, body size still may (Canis latrans) exhibit variability in their playa role in selection of prey. social behavior (Bowen 1981; Harrison Prey selection also may vary among so- 1992; Messier and Barrette 1982) and have cial categories within a predator species as been reported to stalk or lie in wait when a result of differences in behavior or ener- hunting small mammals (Bowyer 1987; getic needs. Male and female mountain li- Wells and Bekoff 1982). Hunting of ungu- ons may encounter different sex and age lates by coyotes, however, typically in- classes of deer at varying frequencies be- volves coursing tactics in which prey are cause of differences in habitat selection, approached, tested, and sometimes pursued timing and amount of movement, or home- over long distances (Bleich 1999; Bowyer range size of these large predators. Ener- 1987; Gese and Grothe 1995). getic needs of male and female mountain Most felids are stalking predators (Ewer lions likely vary because of differences in 1973; Leyhausen 1979) that rely on cover body size or demands of rearing young, but and stealth (Seidensticker 1976; Sunquist 1981) to approach prey closely and then data on this topic are few. rush and pursue an individual over a rela- We compared mortality caused by moun- tively short distance (Bank and Franklin tain lions and coyotes versus that caused by 1998; Elliott et al. 1977; Van OrsdoI1984). automobiles for a single population of mule This form of ambush hunting has been re- deer to examine selection of ungulate prey ported for mountain lions (Puma conco- by predators that differ substantially in lor-Bank and Franklin 1998; Beier et al. body size, hunting style, and reproductive 1995). When prey occur in groups (e.g., status. We predicted that coyotes, a small mule deer, Odocoileus hemionus-Bowyer coursing predator, would be more likely 1984), the stalking technique of felids could than mountain lions, a large stalking hunter, limit their ability to select for young, old, to select mule deer that were younger or in or weakened animals (Schaller 1972). poor physical condition. We also predicted Body size of prey also may influence se- that male and female mountain lions would lection by carnivores (Bekoff et al. 1984; not differ in their selection of prey unless Kunkel et al. 1999). Most ungulates are other factors besides hunting style (e.g., sexually dimorphic, with males substantial- body size or reproductive state) were im- ly larger than females (Ralls 1977; Weck- portant determinants of prey selection. We erly 1998). Additionally, males often pos- predicted that female mountain lions would sess horns or antlers that can increase the kill a greater proportion of young deer than risk of injury to a predator (Hornocker would males and that female mountain li- 1970). Most felids are solitary hunters and ons also would kill a greater proportion of tend to kill species weighing more than half adult female deer than would male moun- their own body weight (Gittleman 1985; tain lions. If body size affected prey selec- Packer 1986). Because male mountain lions tion by coyotes and mountain lions, we also can be >50% larger in body size than fe- males (Dixon 1982), sexual dimorphism predicted that such marked differences in may lead to differences between males and body size would lead to a preponderance of females in ability to kill prey and risk as- small prey items in the diet of coyotes, even sociated with doing so. Furthermore, moun- though these canids often hunt in packs. We tain lions are substantially larger than coy- also predicted that mountain lions with dif- otes; in California, these canids weigh ferent reproductive demands would kill about 9.8-11.2 kg (Hawthorne 1971). Al- mule deer differentially with respect to sex though coyotes can hunt in packs (Bowen and age classes of deer. 464 JOURNAL OF MAMMALOGY Vol. 81, No.2

MATERIALS AND METHODS able collars (Bleich and Pierce 1999); their sex ratio was close to I: I. Twenty-one adult moun- Study area.-Round Valley is located on the tain lions (12 females, 9 males) and 21 offspring east side of the Sierra Nevada in eastern Cali- ( < I year old; 14 male, 7 female) were captured fornia (37°24'N, 118°34'W). Deer inhabit about with the aid of hounds (adults and kittens; n = 90 km2 during November-April, but the area of 38-Davis et al. 1996) or foot snares (n = 4) use varies with snow depth (Kucera 1988). Most during November 1991-May 1996. We weighed mule deer that overwinter in Round Valley mi- mountain lions to the nearest 2.5 kg using a grate in spring to high-elevation ranges in sum- spring scale, and mean weight of adult mountain mer (Kucera 1992; Pierce et al. 1999). A small lions was determined using the Ist recorded proportion of the herd, however, remains on the eastern side of the mountains and is prey for weight for each individual. All adults were fitted with radiocollars. Nine kittens (:56 months old) resident mountain lions and coyotes throughout the year. from 3 litters were captured in natal dens (Bleich Dominant vegetation is characteristic of the et al. 1996). Age of young mountain lions was Great Basin (Storer and Usinger 1968) and in- estimated with weight, pelage characteristics, cludes a mosaic of bitterbrush (Purshia triden- and patterns of tooth eruption (Anderson 1983; tata), sagebrush (Artemisia tridentata), and rab- D. Ashman et al., in litt.). All methods used in bitbrush (Chrysothamnus nauseosum). Patches this research were approved by an Institutional of blackbrush (Coleogyne ramosissima) and Animal Care and Use Committee at the Univer- mormon tea (Ephedra nevadensis) also are in- sity of Alaska-Fairbanks. terspersed. Salix, Rosa, and Betula occidentalis Helicopter surveys were conducted annually occur in riparian areas. Detailed descriptions of in January to determine number of deer in the the study area were provided by Kucera (1992) study area and proportions of adult males, adult and Pierce et al. (1999). females, and young. Transects were flown with Our study began in November 1991 at the end 3 observers and extended across the entire study of a prolonged drought. Annual precipitation area at an elevation where deer tracks in snow during the study was highly variable: the coef- could no longer be seen. ficient of variation of annual precipitation was Deer mortality and predation.-We located 51% during 1983-1998, and annual precipitation mule deer killed by mountain lions and coyotes ranged from 5.3 to 25.2 cm, with 72% occurring during 1991-1998 by back-tracking lions from between November and March. Mean monthly daytime positions, investigating mortality sig- temperatures on the winter range (November- nals from transmitter-equipped deer, locating March) varied from 0°C to 16°C. Estimated mountain lions at night via telemetry, and in- numbers of deer on the winter range increased vestigating lacations of scavenging birds. All gradually over the period of the study from 939 marked deer were monitored daily for mortality (10 deer/km2) in 1991 to 1,913 (21 deer/km2) in signals, and causesof mortality were determined 1997, whereas mean number of mountain lions by examining wounds, tracks, and feces in the decreased from 6.1 in winter 1992-1993 to 3.0 vicinity of carcasses.Additionally, remote pho- in 1996-1997 (Pierce et al., in press). tography (Pierce et al. 1998) facilitated deter- Sampling prey and predators.- Three hundred mination of the predator responsible for a kill. ten mule deer (217 females, 93 males) were cap- Lower incisors and femurs were collected from tured and fitted with radiocollars during winter all carcassesof mule deer for age analysis with or spring 1993-1997. Deer were captured using cementum annuli (Low and Cowan 1963) and Clover traps (n = 9-Clover 1956), drop nets (n for analysis of fat with ether extraction of mar- = 2-Conner et al. 1987), and a net gun fired row in long bones (Neiland 1970). from a helicopter (n = 299-Krausman et al. We collected feces of carnivores opportunis- 1985). Deer were captured on their winter range, tically for analysis of diets. Although many fe- and individuals from groups that already includ- ces for both predators were gathered near loca- ed an animal with a collar were intentionally tions of kills, coyote feces were located through- avoided. Collars were distributed among adult out the study area, especially along roads. Feces males and females in the approximate proportion of mountain lions were numerous at latrines (lo- of their occurrence in the population (1:3). Deer cations used repeatedly for scent marking) and <1 year old (n = 113) were fitted with expand- near resting areas. Feces of mountain lions also May 2000 PIERCE ET AL.-PREDATION BY MOUNTAIN LIONS AND COYOTES 465 were located by hounds while trailing mountain tain lions and coyotes. For comparisons made lions. Identification of food in fecal samples was within the adult age category (e.g., sex and age determined from remains of bone, teeth, and of adults), data from throughout the year were claws and from hair samples examined for color, used because they were not biased by the birth- length, thickness, and medullary characteristics ing season of mule deer. (Big Sky Laboratory, Florence, Montana). Re- Analysis of variance (Neter et al. 1990) was mains identified from carnivore feces were cat- used to test for differences in age of adult deer egorized as mule deer, small animals ( < 15 kg), killed by automobiles, mountain lions, and coy- or other and were summarized as percentage oc- otes. Differences in the percentage of fat in the currence in feces. marrow of adult females killed by vehicles in We used data from deer killed by automobiles March 1993 and 1994 and adult females col- during 1991-1998 to estimate sex, age class, and lected in March of the same years also were physical condition of prey available to predators. evaluated with analyses of variance. We used Use of such animals as a random sample of a multidimensional chi-square analysis (Zar 1984) population has been questioned (O'Gara and to determine if there were differences in cate- Harris 1988) becausedeer in poor condition may gories of age and condition and in sex of mule be more likely to use roadways for paths of trav- deer killed by automobiles, mountain lions, and el through deep snow. This potential problem, coyotes in October-June. Mule deer were cate- however, was not a consideration for our study gorized as young «1 year) or adult and good area. Most deer (55% of 191 deer) killed by au- condition (>50% fat in bone marrow) or poor tomobiles were collected from Highway 395, condition to meet assumptions of chi-square and snow depth rarely was greater than a few analysis (Zar 1984). Use of bone-marrow fat to centimeters in the vicinity of that roadway; deer index condition may be problematic because killed along the roadway were not there to avoid these fat deposits are the last to be used by un- deep snow. To ensure our sample of deer killed gulates (Mech and DelGiudice 1985); therefore, by automobiles was not biased, we tested for an animal that has used most of its body-fat re- differences in age composition of those deer serves and is in poor condition may still have killed during winter (October-April) against some fat in the marrow of their long bones. data obtained from aerial surveys conducted in Measures of body condition based on kidney fat January of each year. and heart fat rarely were available for deer killed In addition to using deer killed by automo- by mountain lions and coyotes because those or- biles as a sample of the deer population, we es- gans often were consumed. When bone-marrow timated proportion of postnatal deer ( <4 months fat in red deer (Cervus elaphus) reached about old) in the population during late summer (July- 50%, kidney fat approached about 25% (Riney September) by fetal rates. Adult females were 1955). Low kidney fat coincides with other in- shot at random annually in March following the dices of malnutrition; therefore, our results as- methods of Kucera (1997). The mean number of sumed that deer with :550% bone-marrow fat fetuses per adult female (n = 86) in 1992-1996 were in poorer condition than those deer with was used to estimate the proportion of postnatal >50% bone-marrow fat. We used multidimen- deer in the population in late summer. Use of sional chi-square analysis to test for differences fetal rates to estimate the available proportion of in age, condition, and sex of mule deer (from all postnatal deer during late summer did not ac- months) killed by automobiles and in social cat- count for mortality and therefore was an over- egories of mountain lions. Social categories of estimate. Thus, our estimate of selection for mountain lions included solitary adult males, postnatal deer by predators was conservative. solitary adult females, adult females with juve- Data analysis.-We used chi-square analysis niles (>6 months old but not independent), and (Zar 1984) to compare the proportion of <1- adult females with kittens (:56 months old- year-old deer killed by automobiles in late sum- Pierce et al. 1998). mer (July-September) with the proportion of Because female mountain lions gave birth to postnatal deer ( <4 months old) expected, based litters in late summer (Bleich et al. 1996), fe- on fetal rates of adult females collected in males with young consistently had a higher pro- March. Consequently, we used only data from portion of young deer available as prey relative October-June to test for prey selection by moun-~ to other social categories. Therefore, we parti-~ 466 JOURNAL OF MAMMALOGY Vol. 81, No.2

TABLE 1.-Proportion of adult (~1 year old) and young «1 year old) mule deer in poor (:s;50% bone-marrow fat) and good (>50% bone-marrow fat) condition killed by automobiles and predators in the eastern Sierra Nevada, California (1991-1998). Proportions were not different among sources of mortality for young deer or adults (P > 0.05).

Deer age and physical condition

tioned our data to test for prey selection by fe- automobiles among young deer or adults male mountain lions with kittens. Using a chi- (Table 1). Additionally, mule deer in poor square test, we compared the proportion of post- condition comprised a small proportion of natal deer killed by female mountain lions with the deer killed by all sources (Table 1). kittens with the proportion of postnatal deer in the population during late summer as estimated Analysis of age, sex, and condition of from fetal rates. mule deer killed by automobiles, mountain lions, and coyotes from October to June in- RESULTS dicated no difference in the sex or condition Analysis of prey remains in carnivore fe- of mule deer for all ages combined (Table ces indicated that mule deer were the pri- 2). Mountain lions, however, killed a great- mary food of mountain lions, occurring in er proportion of young deer than did auto- 73% of feces, whereas remains of mule deer mobiles (P < 0.01; Fig. 1). Coyotes did not occurred in 17% of coyote feces (X2 = select more young (26%) than expected 65.21, d.f. = 1, p < 0.001). Desert cotton- based on the proportion killed by automo- tails (Sylvilagus audubonii) and black-tailed biles (21 %; p = 0.42; Fig. 1). When com- jack rabbits (Lepus califomicus) were the paring sex and age of adult mule deer primary species of small animals in the diet among sources of mortality, mountain lions of mountain lions and coyotes. and coyotes killed more females than did The proportion of young in the popula- automobiles (P < 0.05; Fig. 1), and the tion as determined by aerial surveys (26% mean age of adult deer killed by predators :t 4.4 SD) did not differ from that estimated was greater than that of adult deer killed by from deer killed by automobiles (25% :t automobiles (P < 0.001; Fig. 2). 8.4%) during all years pooled (X2 = 0.16, Mean weight of 8 adult male mountain d.f. = 1, p = 0.69) or during any year (P lions (55 kg ::t:7.7 SD) was >25% the mean ?; 0.19). Additionally, percentage of fat in weight of 11 adult females (40 ::t: 5.1 kg). the bone marrow of mule deer collected in Social categories of mountain lions did not March (78% :t 20%, n = 31) did not differ differ when contrasted with sex or condi- from the percentage observed for adult fe- tion of deer preyed upon by these animals males killed by vehicles in March (64% :t and of deer killed by automobiles, when 19%, n = 6; F = 2.70, d.f. = 1, 36, p = age categories of deer were combined (Ta- 0.10). For deer killed during all months, ble 3). When only adult deer were consid- more young deer than adults in the popu- ered, however, a higher percentage of fe- lation were in poor condition (X2 = 5.6, d.f. male deer were killed by female mountain = 1, p < 0.05; Table 1); however, no se- lions (79% ) than were killed by automo- lection occurred with respect to physical biles (63%; X2 = 6.02, dj= 1, p < 0.05), condition for deer killed by predators or by but female and male (81 %; X2 = 2.70, d.f. May 2000 PIERCE ET AL -PREDAnON BY MOUNTAIN LIONS AND COYOTES 467

TABLE 2.-Proportion of male and female mule deer in poor (~50% bone-marrow fat) or good (>50% bone-marrow fat) condition killed by automobiles or predators in the eastern Sierra Nevada, California, October-June (1991-1998). Data collected from July through September were excluded because of a high proportion of postnatal deer ( <4 months old) in the populations that were under- represented in mortality data. Proportions were not different among sources of mortality for condition or sex (P > 0.05).

Deer physical condition and sex

Source of mortality

Automobiles 81 Mountain lions 78 Coyotes 72

= 1; p = 0.10) mountain lions killed the = 1, p < 0.001); however, solitary females same percentage of female deer. Female did not differ from male mountain lions in mountain lions killed a greater proportion selection of age categories of deer (X2 = of young deer than did males (X2 = 4.81, 1.18, d.f = 1, p = 0.18; Fig. 3). dj = 1, p < 0.05) or automobiles (X2 = Despite the concordance between pro- 31.01, dj = 1, p < 0.001; Fig. 3). Con- portions of young deer killed by automo- sidered separately, solitary females still se- biles during October-April and proportions lected proportionally more young deer than observed in annual surveys, comparison of were killed by automobiles (X2 = 11.12, d.f.

8 = 13.0,P < 0.001 ~ I 22!1 w 7 w O w ..J 6 ~ ~ 1 ... 5 I ..J (97) ~ (19) 0 4 < ~ O 3 w I ~ < (111) 2

1

0 MOUNTAIN LION AUTOMOBILE COYOTE

SOURCEOF MORTALITY FiG. 2.-Mean (:t SE) age of adult mule deer killed by mountain lions, coyotes, and automo- biles in the eastern Sierra Nevada, California, 1991-1998. Ages of deer were determined by analysis of cementum annuli. Adult deer killed by mountain lions and coyotes were older than those killed by automobiles. Sample sizes are shown in parentheses. 468 JOURNAL OF MAMMALOGY Val. 81, Nc

TABLE3.-Proportion of male and female mule deer killed in poor (:550% bone-marrow fat) or good (>50% bone-marrow fat) condition that were killed by automobiles or various social categories of mountain lions in the eastern Sierra Nevada. California (1991-1998). There wa~ no difference (P > 0.05) in the condition or the sex of mule deer killed by mountain lions and those killed by automobiles when all age categories and all months were combined.

Deer physical condition and sex

the proportion of young deer killed by au- (92% ) during late summer compared with tomobiles (15%) and the estimated propor- the proportion of postnatal deer estimated tion of young deer based on fetal rates by fetal rates (X2 = 7.94, d.f = 1, p < (51 %) in late summer indicated that post- 0.01). natal deer were underrepresented in our sample of deer killed by automobiles (X2 = DISCUSSION 21.70, d.f = 1, p < 0.001). Mountain lions Hunting style was not an important factor with kittens still selected postnatal deer in prey selection of mule deer; only moun- tain lions selected young deer (Fig. 1), but both mountain lions and coyotes selected X;=36.7,P6 months old ing predator would not be as selective as a but not independent), and females with kittens coursing predator was rejected because (:56 months). Proportion of young and adult mountain lions were as selective, relative to deer killed by each social category of mountain lion was compared with the proportion available availability of different age and condition as indexed by deer killed by automobiles (0 on categories of prey, as were coyotes. y-axis; % selection = % use -% available). The hypothesis that body size of the May 2000 PIERCE ET AL.-PREDATION BY MOUNTAIN LIONS AND COYOTES 469 predator affects prey selection was support- of males in the deer population suggests a ed for predation by mountain lions and coy- probable increase in the proportion of otes on prey species of different sizes and young animals in that segment of the pop- for predation by mountain lions on adult ulation; therefore, a large proportion of the mule deer. Coyotes had a higher proportion male deer available to mountain lions and of small prey in their diets than did moun- coyotes in Round Valler may have been tain lions, despite the ability of coyotes to younger and not in weakened physical con- hunt in packs and kill large prey (Bowen dition from mating activities. 1981; Bowyer 1987). Collection of feces Predation on both young and adult deer throughout the 6 years of our study helped by both coyotes and mountain lions caused prevent biases associated with population us to reject the hypothesis that body size fluctuations and prey switching in coyotes differences b~tween these 2 predators un- (O'Donoghue et al. 1998; Patterson et al. derlie differences in prey selection because 1998) and biases associatedwith changes in coyotes did not select for young deer (Fig. density and distribution of mountain lions I). Because our results were for October- and their primary prey (Pierce et al. in June, postnatal deer may have been under- press). Coyotes also were more omnivorous represented in predator kills because fewer than mountain lions, which were strict car- remains of young are left and their carcass- nivores, a pattern that has been observed es are more difficult to locate (Johnsingh previously for canids and felids (Bowyer et 1993; Schaller 1967). No deer killed by al. 1983; Litvaitis and Harrison 1989). Fur- coyotes were located during late summer, ther, female mountain lions selected female possibly because coyotes completely con- deer, whereas male mountain lions did not. sumed postnatal deer before we could lo- Selection of female deer by mountain lions cate carcasses. During other studies, coy- in our study was contrary to the findings of otes have preyed heavily on young deer in Hornocker (1970) but was consistent with late summer (Andelt 1985; Bowyer 1987; those of Bleich and Taylor (1998), who re- Litvaitis and Shaw 1980). Coyotes also of- ported that female deer from populations ten hunted in packs in Round Valley (B. M. directly north of Round Valley were killed Pierce, in litt.) and elsewhere (Bowen 1981; by mountain lions in greater proportion ~owyer 1987; Messier and Barrette 1982) than expected based on their frequency in and may benefit energetically by killing the population. Hornocker (1970) reported larger prey (Kruuk 1975; Peterson 1977). selection for male prey and proposed that Female mountain lions rearing offspring male deer and elk (Cervus elaphus) were selected young deer as prey. This outcome weakened during the mating season and may be the result of physiological con- therefore more prone to predation by moun- straints of lactation. During lactation, fe- tain lions. Male ungulates also spatially male mountain lions may need a more con- segregate from females seasonally (Bleich sistant intake of protein than do males. Be- et al. 1997; Bowyer 1984; Bowyer et al. cause protein is stored as muscle, drawing 1996) and may select habitats that place upon protein reserves for too long can af - them at more risk for predation than do fe- fect locomotion (Wannemacher and Cooper males (Bleich et al. 1997; Hornocker 1970). 1970). Young deer may be easier than The proportion of adult male deer on the adults to catch because young animals lack winter range in Round Valley, as indexed experience in escaping predators and may by aerial surveys, increased during 1992- lack stamina (Curio 1976; Schaller 1972); 1997 from 12% to 45% of the adult pOpU- therefore, young deer may provide a small- lation compared with 19% for the popula- er but more consistant source of food. We tion of ungulates in Idaho (Hornocker hypothesize that for lactating females the 1970). Such an increase in the proportion risk of an unsuccessful hunt and the accom- 470 JOURNAL OF MAMMALOGY Vol. 81, No.2 panying drain on protein reserves may demands of reproduction on prey selection overshadow the benefit of killing larger by carnivores is needed to understand pred- prey. Male mountain lions may have lower ator-prey relationships more completely. killing rates, killing larger prey less often and gorging themselves to store fat. Such a ACKNOWLEDGMENTS strategy would allow males to make long- We thank the houndsmen, field technicians, range movements in search of females and and volunteers that helped capture mountain li- in defense of their relatively large territories ons and locate deer killed by mountain lions: I. (Anderson et al. 1992). Davis, V. Davis, I. Ostergard, C. Baker, P. Par- Sexual segregation in both predator and tridge, C.-L. Chetkiewicz, G. Raygorodetsky, prey also may have led to differences in and D. Stoner. We give special thanks to I. We- prey selection with respect to age categories hausen for assistance in capturing mountain li- of deer. Female mountain lions with kittens ons. We also are grateful to the volunteers who helped in daily monitoring of the deer: W. AII- were located at relatively low elevations sup, C. Baker, G. Clarkson, H. Iones, R. Noles, during the late summer compared with oth- D. Noles, N. Partridge, P. Partridge, B. Peters, P. er mountain lions (Pierce et al. 1999). Male Thompson, and the late R. Williamson. We ex- deer segregate from females with young tend special thanks to R. Anthes for his safe fly- during spring and summer (Bowyer 1984; ing. This research was supported by the Califor- Bowyer et al. 1996; Main and Coblentz nia Department of Fish and Game, the National 1996) and consequently may have been en- Rifle Association, Friends of the National Rifle countered at lower frequencies by mountain Association, the Mule Deer Foundation, Safari lions at lower elevations. Club International (Los Angeles, Orange Coun- Our study demonstrated that presence of ty , Sacramento, San Fernando, and San Francis- dependent young was a critical factor af- co Bay Area chapters), the Fish and Game Ad- fecting prey selection by mountain lions. visory Committee of Inyo and Mono Counties, Contrary to our prediction, differences in the California Deer Association, the University of California White Mountain Research Station, hunting style between mountain lions and and the Institute of Arctic Biology at the Uni- coyotes was not important in prey selection; versity of Alaska-Fairbanks. This is a contri- the stalking predator was as selective as the bution from the California Department of Fish coursing predator. Differences in body size and Game Deer Herd Management Plan Irnple- between coyotes and mountain lions may mentation Program and is Professional Paper 15 be important for the selection of prey spe- from the Eastern Sierra Center for Applied Pop- cies but not for the selection of deer from ulation Ecology. October to June. Body size was not an im- portant factor for selection of deer by LITERA TURE CITED mountain lions because solitary male and AcKERMAN, B. B., E G. LINDZEY, AND T. P. HEMKER. female mountain lions both killed young 1984. Cougar food habits in southern Utah. The Journal of Wildlife Management 48:147-155. and adult deer in proportion to their avail- ANDELT, W. E 1985. Behavioral ecology of coyotes in ability. south Texas. Wildlife Monographs 94:1-45. Other investigators (Hornocker 1970; ANDERSON, A. E. 1983. A critical review of literature Mech 1970; Mech and Frenzel 1971) have on puma (Fe/is conc%r). Colorado Division of Wildlife, Special Report 54:1-91. reported similar results for prey selection ANDERSON,A. E., D. C. BOWDEN, AND D. M. KATTNER. among multiple species of prey or by a sin- 1992. The puma on the tJncompahgre Plateau, Col- gle species of predator. Our results, how- orado. Colorado Division of Wildlife, Technical Publication 40:1-116. ever, emphasize the importance of consid- BANK, M. S., AND W. L. FRANKLIN. 1998. Puma (Puma ering reproductive status of the predator conc%r patagonica) feeding observations and at- when attempting to predict how that species tacks on guanacos (Lama guanicoe). M~a1ia 62: 599-605. of predator may affect a prey population. BEIER, P., D. CHOATE, AND R. H. BARRETT. 1995. Further study of the effects of the energy Movement patterns of mountain lions during differ- May 2000 PIERCE ET AL.-PREDATION BY MOUNTAIN LIONS AND COYOTES 471

ent behaviors. Iournal of Mammalogy 76:1056- cal and taxonomic correlates. Oecologia 67:540- 1070. 554. BEKoFF, M., T. I. DANIELS, AND I. L. GITfLEMAN. 1984. HARRISON, D. I. 1992. Social ecology of coyotes in Life history patterns and the comparative social northeastern North America: relationships to dis- ecology of carnivores. Annual Review of Ecology persal, food resources, and human exploitation. pp. and Systematics 15:191-232. 53-72 in Ecology and management of the eastern BLEICH, V. C. 1999. Mountain sheep and coyotes: pat- coyote (A. N. Boer, ed.). Wildlife Research Unit, terns of predator evasion in a mountain ungulate. University of New Brunswick, Fredricton, New Iournal of Mammalogy 80:283-289. Brunswick, Canada. BLEIcH, V. C., R. T. BowYER, AND I. D. WEHAUSEN. HAWTHORNE, V. M. 1971. Coyote movements in Sa- 1997. Sexual segregation in mountain sheep: re- gehen Creek Basin, northeastern California. Califor- sources or predation? Wildlife Monographs 134:1- nia Fish and Game 57:154-161. 50. HoRNOCKER, M. G. 1970. An analysis of mountain lion BLEIcH, V. C., AND B. M. PIERCE. 1999. Expandable predation upon mule deer and elk in the Idaho Prim- and economical radio collars for juvenile mule deer. itive Area. Wildlife Monographs 21:1-39. California Fish and Game 85:56-62. HuGGARD, D. I. 1993. Prey selectivity by wolves in BLEIcH, V. C., B. M. PIERCE, I. L. DAVIS, AND V. L. Banff National Park. I. Prey species. Canadian Jour- DAVIS. 1996. Thermal characteristics of mountain nal of Zoology 71:130-139. lion dens. The Great Basin Naturalist 56:276-278. IOHNSINGH, A. I. T. 1993. Prey selection in three large BLEICH, V. C., AND T. I. TAYLOR. 1998. Survivorship sympatric carnivores in Bandipur. Mammalia 56: and cause-specific mortality in five populations of 517-526. mule deer. The Great Basin Naturalist 58:265-272. KARANTH, K. U., AND M. E. SUNQUIST. 1995. Prey se- BoWEN, D. W. 1981. Variation in coyote social orga- lection by tiger, leopard and dhole in tropical forests. nization: the influence of prey size. Canadian Jour- The Iournal of Animal Ecology 64:439-450. nal of Zoology 59:639-652. KLEIMAN, D. G., AND I. E EISENBERG. 1973. Compar- BowYER, R. T. 1984. Sexual segregation in southern isons of canid and felid social systems from an evo- mule deer. Iournal of Mammalogy 65:410-417. lutionary perspective. Animal Behaviour 21:637- BowYER, R. T. 1987. Coyote group size relative to pre- 659. dation on mule deer. Mammalia 51:515-526. KRAUSMAN, P. R., I. I. HERVERT, AND L. L. ORDWAY. BoWYER, R. T., I. G. K!E, AND V. VAN BALLENBERGHE. 1985. Capturing deer and mountain sheep with a 1996. Sexual segregation in black-tailed deer: effects net-gun. Wildlife Society Bulletin 13:71-73. of scale. The Iournal of Wildlife Management 60: KRUUK, H. 1972. The spotted hyena. The University 10-17. of Chicago Press, Chicago, Illinois. BowYER, R. T., S. A. MCKENNA, AND M. E. SHEA. KRUUK, H. 1975. Functional aspects of social hunting 1983. Seasonal changes in coyote food habits as de- in carnivores. pp. 119-141 in Function and evolu- terrnined by fecal analysis. The American Midland tion in behaviour (G. Baerends, C. Beer, and A. Naturalist 109:266-273. Manning, eds.). Oxford University Press, Oxford, CLOVER, M. R. 1956. Single-gate deer trap. California United Kingdom. Fish and Game 42:199-201. KUCERA, T. E. 1988. Ecology and population dynamics CONNER, M. C., E. C. SOUTIERE, AND R. A. LANCIA. of mule deer in the eastern Sierra Nevada, Califor- 1987. Drop-netting deer: costs and incidence of cap- nia. Ph.D. dissertation, University of California, ture myopathy. Wildlife Society Bulletin 15:434- Berkeley. 438. KUCERA, T. E. 1992. Influences of sex and weather on CURIO, E. 1976. Ethology of predation. Springer-Ver- migration of mule deer in California. The Great Ba- lag, Berlin, Germany. sin Naturalist 52: 122-130. DAVIS, I. L., ET AL. 1996. A device to safely remove KUCERA, T. E. 1997. Fecal indicators, diet, and popu- immobilized mountain lions from trees and cliffs. lation parameters in mule deer. The Iournal of Wild- Wildlife Society Bulletin 24:537-539. life Management 61:550-560. DIXON, K. R. 1982. Mountain lion. pp. 711-727 in KUNKEL, K. E., T. K. RUTH, D. H. PLETSCHER,AND M. Wild mammals of North America (I. A. Chapman G. HoRNOCKER. 1999. Winter prey selection by and G. A. Feldhamer, eds.). The Iohns Ropkins Uni- wolves and cougars in and near Glacier National versity Press, Baltimore, Maryland. Park. The Iournal of Wildlife Management 63:901- ELLIOT, I. P., I. McT. COWAN, AND C. S. ROLLING. 1977. 910. Prey capture by the African lion. Canadian Iournal KuYT, E. 1972. Food habits of wolves on barren- of Zoology 55:1811-1828. ground caribou range. Canadian Wildlife Service EsTES, R. D., AND I. GODDARD. 1967. Prey selection Report 21:1-35. and hunting behavior of the African wild dog. The LEYHAUSEN,P. 1979. Cat behavior: the predatory and Iournal of Wildlife Management 31:52-70. social behavior of domestic and wild cats. Garland EWER, R. E 1973. The carnivores. Cornell University STPM Press, New York. Press, Ithaca, New York. LITVArrIS, I. A., AND D. I. HARRISON. 1989. Bobcat- GESE, E. M., AND S. GROTHE. 1995. Analysis of coyote coyote niche relationships during a period of coyote predation on deer and elk during winter in Yellow- population increase. Canadian Iournal of Zoology stone National Park, Wyoming. The American Mid- 67:1180-1188. land Naturalist 133:36-43. LITVArrIS, I. A., AND I. H. SHAw. 1980. Coyote move- GITfLEMAN, I. L. 1985. Carnivore body size: ecologi- ments, habitat use, and food habits in southwestern 472 JOURNAL OF MAMMALOGY Vol. 81, No.2

oklahoma. The Journal of Wildlife Management 44: land-tenure system regulate population size? Ecolo- 62-68. gy. Low, W. A., AND I. M. COWAN. 1963. Age determi- PIERCE, B. M., V. C. BLEICH, C.-L. B. CHETKIEWICZ, nation of deer by annular structure of dental cemen- AND J. D. WEHAUSEN. 1998. Timing of feeding bouts tum. The Journal Wildlife Management 27:466-471. of mountain lions. Journal of Mammalogy 79:222- MAIN, M. B., AND B. E. COBLEN1Z. 1996. Sexual seg- 226. regation in Rocky Mountain mule deer. The Journal PIERCE, B. M., V. C. BLEICH, J. D. WEHAUSEN, AND R. Wildlife Management 60:497-507. T. BOWYER. 1999. Migratory patterns of mountain MCLAREN, B. E., AND R. 0. PETERSON.1994. Wolves, lions: implications for social regulation and conser- moose, and tree rings on Isle Royale. Science 266: vation. Journal of Mammalogy 80:986-992. PIMLOTf, D. H., J. A SHANNON, AND G. B. KOLENSKY. 1555-1558. 1969. The ecology of the timber wolf in Algonquin MECH, L. D. 1970. The wolf: ecology and behavior of Park. Ontario Department of Lands and Forests, an endangered species. Natural History Press, New Wildlife Research Report 87:1-92. York. RALLS, K. 1977. Sexual dimorphism in mammals: avi- MECH, L. D., AND G. D. DELGIUDICE. 1985. Limitations an models and unanswered questions. The American of the marrow-fat technique as an indicator of body Naturalist 111:917-938. condition. Wildlife Society Bulletin 13:204-206. RiNEY, T. 1955. Evaluating condition of free-ranging MECH, L. D., AND L. D. FRENZEL. 1971. Ecological red deer (Cervus elaphus), with special reference to studies of the timber wolf in north-eastern Minne- New Zealand. New Zealand Journal of Science and sota. United States Forest Service Research Paper Technology Section B 36:429-463. NC 52:1-62. Ross, P. I., AND M. G. JALKOTZY. 1996. Cougar pre- MEsSIER, E, AND C. BARRETfE. 1982. The social system dation on moose in south-western Alberta. Alces 32: of the coyote ( Canis latrans) in a forested habitat. 1-8. Canadian Journal of Zoology 60: 1743-1753. SCHALLER, G. B. 1967. The deer and the tiger: a study MuRIE, A. 1944. The wolves of Mount McKinley. Fau- of wildlife in India. The University of Chicago na of the National Parks of the United States, Fauna Press, Chicago, Illinois. Series 5:1-238. SCHALLER, G. B. 1972. The Serengeti lion. The Uni- NEILAND, K. A. 1970. Weight of dried marrow as an versity of Chicago Press, Chicago, Illinois. indicator of fat in caribou femurs. The Journal of SEIDENSTICKER,J. 1976. On the ecological separation Wildlife Management 34:904-907. between tigers and leopards. Biotropica 8:225-234. NETER, J., W. WASSERMAN, AND M. H. KUTNER. 1990. SPALDJNG,D. J., AND J. LESOWSKI. 1971. Winter food Applied linear statistical models. Richard D. Irwin, of the cougar in south-central British Columbia. The Homewood, Illinois. Journal of Wildlife Management 35:378-381. O'DoNOGHUE, M., S. BOU11N, C. J. KREBS, G. ZULETA, STORER,T. I.,. AND R. L. USJNGER.1968. Sierra Nevada D. L. MURRAY, AND E. J. HopER. 1998. Functional natural history. University of California Press, responses of coyotes and lynx to the snowshoe hare Berkeley. SUNQUIST, M. E. 1981. The social organization of ti- cycle. Ecology 79:1193-1208. gers (Panthera tigris) in Royal Chitwan National O'GARA, B. W., AND R. B. HARRIS. 1988. Age and Park, Nepal. Smithsonian Contributions in Zoology condition of deer killed by predators and automo- 336:1-98. biles. The Journal of Wildlife Management 52:316- TAYLOR, R. J. 1984. Predation. Chapman and Hall, 320. New York. PACKER, C. 1986. The ecology of sociality in felids. V AN ORSDOL, K. G. 1984. Foraging behavior and hunt- pp. 429-451 in Ecological aspects of social evolu- ing success of lions in Queen Elizabeth National tion: birds and mammals (D. I Rubenstein and R. Park, Uganda. African Journal of Ecology 22:79- W. Wrangham, eds.). Princeton University Press, 99. Princeton, New Jersey. W ANNEMACHER,R. W., JR., AND W. K. COOPER. 1970. PACKER, C., D. SCHEEL, AND A. E. PuSEY. 1990. Why Relationship between protein metabolism in muscle lions form groups: food is not enough. The Ameri- tissue and the concept of "protein reserves. " pp. can Naturalist 136:1-19. 121-138 in Protein metabolism and biological func- PATfERSON, B. R., L. K. BENJAMIN, AND E MESSIER. tion (C. P. Bianchi and R. Hilf, eds.). Rutgers Uni- 1998. Prey switching and feeding habits of eastern versity Press, New Brunswick, New Jersey. coyotes in relation to snowshoe hare and white- WECKERLY, E W. 1998. Sexual-size dimorphism: influ- tailed deer densities. Canadian Journal of Zoology ence of mass and mating systems in the most di- 76:1885-1897. morphic mammals. Journal of Mammalogy 79:33- PETERSON,R. 0. 1977. Wolf ecology and prey rela- 52. tionships on Isle Royale. United States National WELLS, M. C., AND M. BEKOFF. 1982. Predation by Park Service Scientific Monograph 11:1-210. wild coyotes: behavioral and ecological analyses. Journal of Mammalogy 63:118-127. PETERSON,R. 0., N. J. THOMAS, J. M. THURBER, J. A. ZAR, J. H. 1984. Biostatistical analysis. 2nd ed. Pren- VucEncH, AND T. A. WArm. 1998. Population lim- tice-Hall, Englewood Cliffs, New Jersey. itation and the wolves of Isle Royale. Journal of Mammalogy 79:828-841. Submitted 20 Apri11999. Accepted 4 September 1999. PIERCE, B. M., V. C. BLEICH, AND R. T. BowYER. In press. Social organization of mountain lions: does a Associate Editor was John A. Litvaitis. American Society of Mammalogists, 1999-2000 Standing Committees

Animal Care and Use: A. Krevitz, Chair; G. W. Barrett, J. E. Childs, J. W. Dragoo, T. H. Horton, S. Jenkins, M. Johnson, J. Kenagy, C. Kodnai, T. H. Kunz, T. J. McIntyre, D. K. Odell, P. Paquet, A. J. Pinter,W. E. Sera, R. S. Sikes, and T. L. Yates. Archives: H. H. Genoways, Historian; A. L. Gardner, Archivist. Checklist: D. E. Wilson, Chair; C. G. Anderson, R. L. Brownell, M. D. Carleton, F. Dieterlen, A. L. Gardner, C. P. Groves, P. Grubb, L. R. Heaney, R. S. Hoffmann, M. E. Holden, R. Hutterer, J. G. Mead, G. G. Musser, B. D. Patterson, J. L. Patton, D. A. Reeder, D. A. Schlitter, H. Shoshani, N. B. Simmons, R. W. Thorington, Jr., C. A. Woods, and W. C. Wozencraft. Conservation of Land Mammals: S. R. Sheffield, Chair; B. J. Bergstrom, B. R. Blood, T. J. Doonan, K. R. Foresman, T. Gehring, J. Goheen, R. Hoyt, R. Kays, A. Krevitz, T. E. Lee, K. LoGiudice, C. A. Long, R. Manson, J. E. Moyers, v. Naples, G. Nordquist, B. Pierce, D. Rosenblatt, J. Rachlow, L.Randa, J. Ray, R. Reynolds, andJ. Sandel. Development: S. Anderson, Chair; R. J. Baker, E. C. Bimey, J. R. Choate, H. H. Genoways, S. B. George, A. V. Linzey, T. J. McIntyre, D. K. Odell, D. J. Schmidly, H. D. Smith, B. R. Stein, B. J. Verts, M. R. Willig, D. E. Wilson and T. L. Yates. Education and Graduate Students: S. C. Loeb, Chair; B. K. Clark, T. J. Doonan, E. J. Finck, C. Rice, H. E. Hoekstra, G. A. Kaufman, J. H. Kruper, E. A. Lacey, R. H. Manson, B. R. McMillan, T. E. Nupp, I. M. Ortega, J. D. Peles, L. L. Peppers, C. E. Rebar, W. E. Sera, P. Stapp, R. K. Swihart, M. G. Tannenbaum, G. Tumer-Erfort, and G. M. Wilson. Grants-in-Aid: R. D. Bradley, Chair; L. N. Carraway, B. J. Danielson, J. W. Demastes, T. L. Derting, M. R. Gannon, S, H. Jenkins, F. Keesing, G. J. Kenagy, M. H. Kesner, G. R. Michener, J. A. Randall, W. E. Sera, and P. Sudman. Grinnell Award Committee: B. R. Stein, Chair; G. N. Cameron, B. S. Clark, T. L. Derting, D. J. Hafner, E. Homer, P. Myers, L. Olson, R. S. Ostfeld, B. D. Patterson, and R. W. Thorington, Jr. Honoraria: C. E. Rebar, Chair; V. Hayssen, T. H. Horton, F. Keesing, L. L. Janecek, S. Loeb, J. F. Merritt, D. 0. Ribble, D. S. Rogers, and R. K. Swihart. Honorary Membership: J. H. Brown, Chair; R. J. Baker, E. C. Bimey, A. V. Linzey, andJ. L. Patton. Human Diversity: F. A. Smith, Chair; D. M. Kaufman, D. W. Kaufman, F. Keesing, K. Lyons, C. C. Maguire, B. McMil- Ian, G. R. Michener, I. M. Ortega, M. H. Schadler, M. C. Sullivan, R. W. Thorington, Jr., G. Tumer-Erfort, and B. C. Woods. Index: M. D. Engstrom, Chair; J. L. Eger, V. Hayssen, B. K. Lim, and P. D. Ross. Informatics: S. B. McLaren, Chair; P. V. August, C. L. Frank, W. L. Gannon, C. S. Hood, M. E. Hopton, L. L. Janecek, K. M. Kramer, R. R. Monk, R. Pfau, M. H. Schadler, R. S. Sikes, N. A. Slade, H. B. Smith, B. R. Stein, M. Stokes, P. D. Sudman, and R. W. Thorington, Jr. International Relations: R. A. Medellin, Chair; C. Chimimba, J. A. Cook, L. Emmons, B. Fox, M. R. Gannon, G. Glass, H. Hoekstra, C. Iudica, G. J. Kenagy, T. E. Lacher, E. Lessa, W. Z. Lidicker, Jr., A. Linzey, I. M. Ortega, J. L. Patton, L. Ruedas, J. A. Salazar-Bravo, D. A. Schlitter, A. T. Smith, I. Tiemann, and T. L. Yates. Jackson Award: J. 0. Whitaker, Jr., Chair; R. J. Baker, E. C. Bimey, M. D. Carleton, C. A. Jones, K. S. Kilbum, J. A. Lackey, and B. J. Verts. Latin American Fellowship: B. D. Patterson,Chair; J. K. Braun, A. Castro-Campillo, G. Ceballos, R. A. Medellin, and D. E. Wilson. Legislation and Regulations: J. H. Shaw, Chair; P. K. Anderson, W. L. Gannon, A. Krevitz, T. J. McIntyre, J. D. Peles, C. E. Rebar, S. R. Sheffield, and M. Smolen. Mammallmages Library: R. S. Sikes, Chair; E. J. Finck, Business Manager; M. R. Gannon, J. H. Harris, M. E. Hopton, K. S. Kilbum, K. Kramer, J. A. Lackey, K. Lyons, C. G. Mahan, C. M. Moore, D. W. Moore, and J. 0. Whitaker, Jr. Marine Mammals: P. K. Anderson, Chair; A. Aguayo L., D. J. Boness, R. L. Brownell, V. M. da Silva, S. D. Dawson, D. P. Domning, P. D. Goley, M. Johnson, K. W. Kenyon, L. F. Lowry, C. D. Marshall, T. J. McIntyre, D. K. Odell, T. J. O'Shea, C. A. Potter, M. Smolen, M. Sommer, andJ. A. Thomas. Membership: L. N. Carraway, Chair; B. S. Argogast, F. A. Cervantes, C. T. Chimimba, E. J. Finck, N. Formozov, J. J. Jacquot, C. E. Rebar, T. D. Reynolds, H. R. Roberts, M. Silva, B. J. Verts, and G. M. Wilson. , Merriam Award: G. N. Cameron, Chair; J. H. Brown, M. D. Engstrom, L. R. Heaney, R. S. Ostfeld, C. E. Rebar, B. Riddle, and B. A. Wunder. Nomenclature: A. L. Gardner, Chair; H. N. Bryant, D. P. Domning, J. L. Eger, M. S. Hafner, C. Jones, C. A. Jones, J. L. Patton, and D. E. Wilson. Planning and Finance: T. Kunz, Chair; A. V. Linzey, R. S. Ostfeld, B. D. Patterson, H. D. Smith, B. R. Stein, R. M. Timm, D. E. Wilson, and T. L. Yates. Program: W. P. Smith, Chair; R. E. Barry, J. A. Cranford, K. R. Foresman, K. S. Kilbum, E. Lacey, J. Litvaitis, C. C. Magui.re, R. Manson, K. McBee, B. D. Odell, D. Post, N. Solomon, and S. D. West. Public Education: P. Vaughn, Chair; A. M. Ditto, T. J. Doonan, S. Y. Erdle, R. Gettinger, D. A. Ginnett, S. Gummer, B. Hart, M. Hopton, A. Joachim, D. A. Kelt, R. Kunkel, E. Muths, C. Ruedebusch, A. Stribling, and T. Tomasi. Publications Committee: D. E. Wilson, Chair; E. Anderson, R. E. Barry, T. L. Best, L. N. Carraway, J. A. Cook, M. D. Engstrom, M. J. Hamilton, V. Hayssen, D. M. Leslie, Jr., S. B. McLaren, J. F. Merritt, S. Lariviere, J. A. Litvaitis, B. Riddle, C. A. Schmidt, R. M. Timm, T. E. Tomasi, C. R. Tumlison, K. T. Wilkins, and M. R. Willig. Resolutions: E. J. Heske, Chair; R. J. Baker, J. A. Cook, J. K. Prey, D. M. Kaufman, T. J. Mclntyre, J. H. Shaw, S. R. Sheffield, W. T. Stanley, and R. W. Thorington, Jr. Systematic Collections: W. L. Gannon, Chair; S. T. Alvarez-Casteiieda, G. D. Baumgardner, M. A. Bogan, J. R. Choate, J. A. Cook, R. C. Dowler, M. D. Engstrom, D. J. Hafner, M. S. Hafner, T. Holmes, S. R. Hoofer, N. D. Moncrief, E. A. Rickart, D. S. Rogers, J. A. Salazar-Bravo, N. B. Simmons, W. T. Stanley, andp. X. Villablanca. ad hoc Committee Review: K. McBee, Chair; J. R. Choate, E. J. Heske, C. Lopez-G. and T. A. Spradling. ad hocOfficers Manual: H. H. Genoways, Chair; R. J. Baker, T. L. Best, J. R. Choate, L. L. Janecek, G. Kaufman, H. D. Smith, D. Smith, and T. L. Yates. TheAmerican Society of Mammalogistsis dedicatedto promotinginterest in mammalsthroughout theworld through research, education, and communication among scientists and the generalpub- lic. The ASMaccomplishes this throughannual meetings, international conferences, committee activities,research and educational awards, and publication of scientificand non-technicalmate- rials,including the Journalof Mammalogy.All responsibletypes of researchon mammalsare en- couragedby the ASM. For additionalinformation on the ASM, visit our home page at www.mammalsociety.organd www.mammalogy.org Officers

0. JamesReichman, President LauraL. Janecek,Recording Secretary ThomasH. Kunz,President-Elect H. DuaneSmith, Se~tary-Treasurer BruceD. Patterson,Vice-President DavidM. Leslie,Jr., Journal Editor

Living Past Presidents DonaldF. Hoffmeister Jamess. Findley James H. Brown JamesN. Layne J. MaryTaylor JamesL. Patton SydneyAnderson HughH. Genoways RobertJ. Baker WilliamZ. Lidicker,Jr. DonE. Wilson AliciaV. Linzey RobertS. Hoffmann ElmerC. Birney

Other Elected Directors

1997-2000 1998.2001 1999.2002 Glennis A. Kaufman Barbara H. Blake Lawrence R. Heaney SusanC. Loeb Jorge Salazar-Bravo Karen McBee Richard S. Ostfeld Guy N. Cameron Chris C. Maguire Jay F. Storz Edward J. Heske Felisa A. Smith Richard W. Thorington, Jr. Suzanne B. McLaren Nancy G. Solomon Michael R. Willig Rodrigo A. Medellin Ginny Turner-Erfort

Trustees

Sydney Anderson (2001) Jerry R. Choale (2002) Terry L. Vales (2000)

MEMBERSHIPSAND SUBSCRIPTIONS

Personsactively interested in mammalogymay become members of the ASMupon nomination, paymentof dues,and election. Applications for membershipand informationabout the ASMand the disciplineof mammalogymay be obtainedfrom the Secretary-Treasurer. The Journalof Mammalogyis sentfree to all membersof the AmericanSociety of Mammalogistsnot in arrears for dues.Dues are $35.00 per year. Subscription rates are $170.00per year. Defective copies or copieslost in shipmentwill be replacedif a writtenrequest is receivedwithin 3 monthsof issue.

Nominationsand dues for membership,dues for subscriptions,changes of address, and requests for missingnumbers or numberslost in shipmentshould be addressedto the Journalof Mammalogy, Allen Marketing & Management,810 East 1Oth Street, P.O. Box 1897, Lawrence, KS 66044-8897; 785-843-1235;FAX 785-843-1274.