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Francis J. Singer

Publications and source records attributed to Francis J. Singer.

14 recordsLinked to original sources

Demography of the Pryor Mountain wild horses, 1993-2007

Wild horses (Equus caballus) at Pryor Mountain were studied by direct observation from 1993 through 2007. All horses present were individually identifiable on the basis of coat coloration, head and leg markings, gender, and band associations. Of the 609 horses either present prior to foaling in 1993 or born since, ages were precisely known for 491 (observed as a foal). Ages for 52 horses were estimated through tooth eruption and wear patterns, and for the remaining 66 horses through body size, morphology, and anecdotal evidence concerning when they were present on the range. At varying intensities, never less than 30 days per year, all horses were inventoried and their band associations noted. Foals were paired with dams based on observations of attachment during the early days and weeks of life. Year of death was determined by identification of the carcass where possible. In the absence of finding a carcass, an animal that was not observed for 2 years was considered to have died in the year that it went missing. Animals that were removed from the herd and mares that were part of a contraception study were excluded from calculations of survival and foaling rates, respectively, as appropriate. The average prefoaling population over the 15 years of the study was 148.8 animals (range = 120-187), and the annual foal crop averaged 32.1 (range = 23-40). Large removals (19-60 animals) in four years helped maintain the herd at this level; apparent growth rate (calculated as though removals had not occurred) was 9.6 percent annually (? = 1.096, range = 0.977-1.220). This annual growth rate is relatively low compared to that for many western horse herds, at least in part because of a decline in foal survival. Sex ratio of the foal crop varied widely among years, but pooled across years did not differ from 50:50. Sex ratio in the herd changed mostly as a result of removals. The average age of both males and females in the herd increased during the course of the study. Annual survival of males did not differ from that of females, nor did gender affect annual survival of foals. Pooled across years, ages, and sexes, the annual survival rate was 0.899. Annual foal survival rate was 0.697 and declined through time, with a tendency toward recovery in 2005-2007. Foal survival was higher in larger bands, but did not differ between foals born to primiparous and multiparous mares. A few 2-year-old mares produced foals; foaling rate (excluding contracepted mares and foals they produced) increased through age 10, remained high through age 15, and declined thereafter. Overall foaling rate for mares =3 years of age was 0.576 foals per mare, with no apparent trend during the period of our study. Foaling rate in years following gathers was somewhat lower than in other years. There was a positive relation between foaling rate and band size. Primiparous mares were somewhat less likely to foal in the following year than were multiparous mares. Most stallions that acquired a harem did so at age 5 or 6, and the average age of harem stallions increased during our study. Most harems had 1-3 mares =2 years of age, but harem size varied with age of the stallion, increasing through about age 11 and declining thereafter. About 6 percent of bands had a satellite stallion (=5 years of age), but the mean number of mares did not differ between single- and multistallion bands. Most stallions left their natal band at age 2 or 3, but 17 percent remained with their natal band until age 4 or 5. Foal survival rate was positively related to precipitation, suggesting a possible link to forage production and availability mediated through mare fitness. There also was evidence for density-dependent population regulation, as both population growth rate and survival rate were negatively correlated with population size from the previous year. These and other factors were not sufficient to stabilize the population during our period of study, however, as evidenced by the necess

Scientific Investigations Report

Aspen Ecology in Rocky Mountain National Park: Age Distribution, Genetics, and the Effects of Elk Herbivory

Lack of recruitment and canopy replacement of aspen (Populus tremuloides) stands that grow on the edges of grasslands on the low-elevation elk (Cervus elaphus) winter range of Rocky Mountain National Park (RMNP) in Colorado have been a cause of concern for more than 70 years. We used a combination of traditional dendrochronology and genetic techniques as well as measuring the characteristics of regenerating and nonregenerating stands on the elk winter range to determine when and under what conditions and estimated elk densities these stands established and through what mechanisms they may regenerate. The period from 1975 to 1995 at low elevation on the east side had 80-95 percent fewer aspen stems than would be expected based on the trend from 1855 through 1965. The age structure of aspen in the park indicates that the interacting effects of fires, elk population changes, and livestock grazing had more-or-less consistent effects on aspen from 1855 to 1965. The lack of a significant change in aspen numbers in recent decades in the higher elevation and west side parts of the park supports the idea that the extensive effects of elk browsing have been more important in reducing aspen numbers than other factors. The genetic variation of aspen populations in RMNP is high at the molecular level. We expected to find that most patches of aspen in the park were composed of a single clone of genetically identical trees, but in fact just 7 percent of measured aspen patches consisted of a single clone. A large frequency of polyploid (triploid and tetraploid) genotypes were found on the low elevation, east-side elk winter range. Nonregenerating aspen stands on the winter range had greater annual offtake, shorter saplings, and lower density of mid-height (1.5-2.5 m) saplings than regenerating stands. Overwinter elk browsing, however, did not appear to inhibit the leader length of aspen saplings. The winter range aspen stands of RMNP appear to be highly resilient in the face of very intense herbivory by elk and harsh environmental conditions. Conservation efforts through fencing protection and decreased elk browsing pressure are already being planned as part of the park's new elk management plan. If these efforts are undertaken, conditions that encourage stem recruitment to the tree canopy will likely result and the continued survival of these aspen stands will be enhanced.

Open-File Report

Effects of elk herbivory on vegetation and nitrogen processes

We used 35-year and 4-year ungulate exclosures to determine the effects of elk ( Cervus elaphus ) herbivory on above-ground and below-ground production and soil fertility on the elk winter range in Rocky Mountain National Park (RMNP), Colorado, USA. We used paired grazed and ungrazed plots to evaluate ungulate herbivory effects in short and tall willow ( Salix spp.), aspen ( Populus spp.), and upland grass/shrub vegetation associations. We measured nitrogen (N) fluxes (litter deposition, fecal and urinary deposition from elk, movements of N by elk, N mineralization, soil N availability, elk consumption rates) within the elk winter, above-ground and below-ground N pools (herbaceous, shrub and root biomass, %N in plants, roots, and soil), and N fluxes on and off the elk winter range (seasonal movement of N by elk). Nitrogen mineralization and soil nitrate (NO 3 ) pools were reduced in the short willow community ( P = 0.07 and 0.10, respectively; n = 4 sites) in grazed plots, but not in the upland grass/shrub community or tall willow sites ( P >0.10). Annual growth of willows was reduced by 98% in grazed plots, relative to 35-year exclosures, and 66% relative to 4-year exclosures. Thus, height, canopy size, and litter biomass of willows were reduced, and N yield of willows was 64% less in grazed plots. We evaluated movement of N by elk among 6 major vegetation associations and found that elk grazed more and bedded less in willow vegetation association compared to mixed conifer, mesic meadow, and grassland/shrub associations ( P = 0.014, 0.001, and 0.026, respectively), suggesting that elk herbivory and movement led to a net loss of N in the willow vegetation association. Elk spent less total time in willows than mesic meadow association, yet they consumed large amounts of willow plant biomass. We recommend management of elk numbers and elk herbivory that takes into consideration impacts to N process function, as negative effects from current levels of herbivory were observed in ≥1 of 3 vegetation associations studied.

Colorado

Annual Report for 2003 Wild Horse Research and Field Activities

As stated in the Wild Horse Fertility Control Field Trial Plan, the Bureau of Land Management (BLM) has an immediate need for a safe, effective contraceptive agent to assist in the management of the large number of wild horses on western rangelands. The BLM and the U.S. Geological Survey-Biological Resources Discipline (USGS/BRD) are testing the immunocontraceptive agent Porcine Zonae Pellucida (PZP) in field trials with three free-roaming herds of western wild horses. Extensive research has already been conducted on the safety, efficacy, and duration of PZP applications in both domestic and feral horses on eastern barrier islands and in some select trials I with wild horses in Nevada managed by the BLM. However, significant questions remain concerning the effects of I PZP application at the population level in the wild, as well as effects at the individual level on behavior, social structure, and harem dynamics of free-ranging animals. These questions are best answered with field trials on wild horse herds under a tight research protocol. The ultimate goal is to provide the BLM with the protocols and information necessary to begin using fertility control to regulate population growth rates in wild horse herds on a broader scale. Fertility control is intended to assist the conventional capture, removal, and adoption process as a I means of controlling excess numbers of wild horses and burros, and to greatly reduce the adoption costs and numbers of animals handled. Fertility control is not intended to totally replace the removal and adoption process. The USGSIBRD began assisting the BLM with field trials of immunocontraceptive fertility control of wild horses in early 2001. The first PZP treatments were applied during gathers at the Pryor Mountain Wild Horse Range in September 2001, and the Little Book Cliffs Wild Horse Range, Colorado, in July 2002. At those gathers, 5 horses were treated in the Pryor Mountain WHR, and 23 were treated in the Little Book Cliffs WHR with PZP. These initial treatments were followed by booster injections in 2002. The second injection is required in order to raise, and I maintain, the titer levels of mares high enough to be considered contracepted. By the end of 2002, 13 horses on the Pryor Mountain WHR had received both injections, as had 11 horses in the Little Book Cliffs WHR. In 2003, intensive research efforts were carried out by the USGSIBRD at three field locations; Pryor Mountain WHR, Little I Book Cliffs WHR, and McCullough Peaks Wild Horse Management Area. The work at these sites during this I calendar year included treatment of wild horse mares with PZP in the Pryor herd and Little Book Cliffs herd, development and implementation of behavioral research to investigate potential affects of PZP treatment, continued tracking of demography and foal production in all three herds, and early phases of investigating aerial population estimation survey techniques. Detailed descriptions of these research topics can be found in the Wild Horse and Burro Management Strategic Research Plan and the Wild Horse Fertility Control Field Trial Plan. Field work in 2003 was conducted by USGSIBRD and BLM staff with the assistance of many dedicated individuals. See I Acknowledgments for more details. This report is meant to highlight the activities of the 2003 field season, as well as to provide a general overview of the data collected. More in-depth data analysis will be conducted following the conclusion of each I phase of the research project, and in many cases will not be possible until several seasons of data are collected.

Open-File Report

An animal location-based habitat suitability model for bighorn sheep and wild horses in Bighorn Canyon National Recreation Area and the Pryor Mountain Wild Horse Range, Montana, and Wyoming

The purpose of this habitat suitability model is to provide a tool that will help managers and researchers better manage bighorn sheep and wild horses in the Bighorn Canyon National Recreation Area (BICA) and Pryor Mountain Wild Horse Range (PMWHR). A concern in the management of the Pryor Mountain wild horse population is whether or not the wild horses compete with bighorn sheep for available forage or available space. Two studies have been conducted that have shown no obvious, convincing competition between the two species. A study of diets and habitat-use of both species revealed substantial diet overlap only during some seasons, but there were considerable spatial and habitat separations between wild horses and bighorns during all seasons (Kissell and others, 1996). This empirical data was then used in a modeling exercise that predicted that neither the current (about 160 horses at the time of the analysis) nor larger numbers of wild horses on the area (e.g., about 200 horses) would result in reduced numbers or condition of bighorn sheep (Coughenour 1999). But competition is a very complex biological process to document. Bighorns might have already been spatially avoiding wild horses when these studies were conducted. A second concern for managers is that earlier studies suggest both species are not using many areas of the range that appear to be suitable (Gudorf and others, 1996; Kissell and others, 1996). A primary goal for the management of both species is to increase their numbers for purposes of genetic conservation and viability. The bighorn sheep population declined during the mid-1990’s from a peak of about 211 animals to ~ 100 animals at present. Absolute minimum goals for genetic viability in the bighorn sheep herd (genetic effective population size of N >50) suggest at least 150 animals should be present, while studies of persistence suggest populations of 250+ are e more likely to recover rapidly and persist should the population experience an epizootic die-off (Singer and others, 2001). Since all bighorn sheep populations are potentially vulnerable to disease epizootics, managing for larger populations of 200–300 animals appears to increase the potential for long-term persistence (Berger, 1990; Singer and others, 2001). Wild horses are not prone to rapid disease die-offs. However, minimum goals for genetic viability in the Pryor Mountain wild horses ( Ne > 50) require that at least 160 animals be present on the range (Singer and others, 2000). Since the Ne > 50 goal is set for the breeding of domestic animals, and since the vagaries of drought, severe winters, predation, and other stochastic events cause stress in wild animals, larger goals for Ne (e.g. Ne > 100) for wild horses are even more desirable (USDI, BLM, 1999; Gross, 2000). Expanding the area of the wild horse range is one option, but the prospects for expanding the range do not appear to be great (L. Coates-Markle, BLM, oral comm.). A second option would be to increase the amount of useable habitat for horses on the existing range. One goal of this modeling effort was to use GIS-based habitat analyses to determine the reason wild horses are not using some areas of the range, and to explore the potential for making some of these areas useable. The National Park Service (NPS) has shown considerable interest in management actions within BICA that will increase the range, useable habitat, and population size of bighorn sheep. There has also been interest expressed by the Bureau of Land Management (BLM) and wild horse advocates to improve the useable habitat for wild horses and to possibly increase the size of the horse range.

Report

Bighorn sheep ( Ovis canadensis ) survivorship and habitat studies in Bighorn Canyon National Recreation Area and surrounding lands, Wyoming and Montana, 2000–2003

In the 1850s, bighorn sheep (Ovis canadensis) were numerous and distributed throughout the Bighorn and Pryor Mountains of Montana and Wyoming. After European settlement, bighorn sheep populations declined, and local extinctions occurred in much of their historic range in the western United States. The current bighorn sheep population of Bighorn Canyon National Recreation Area (BICA) is the product of several reintroductions into BICA and surrounding lands. Following a release in 1973 and growth rates near maximum potential of 19.8% per year, the population grew to an estimated peak population of about 211 animals in 1993 and 1994 (Kissell and others, 1996). Recent counts indicate the bighorn sheep population has declined. Kissell and others (1996) reported that the population began to decline rapidly in 1995 and 1996. He noted low ewe:lamb ratios during the decline phase. Bighorn sheep numbers declined to the lowest minimum viable population size of 100 animals recommended by several bighorn sheep experts (Bailey, 1990; Berger, 1990; Smith and others, 1991). National Park Service (NPS) and Bureau of Land Management (BLM) managers were concerned about the decline and requested a study of its causes. In 2000, the U.S. Geological Survey- Biological Resources Division (USGS-BRD) received funding to start a 3-year study of survivorship, condition, and population growth rate of the BICA bighorn sheep population. Several possibilities exist for the bighorn sheep decline. The herd may have experienced a rapid population expansion, followed by a decline to stability at a lower long-term carrying capacity. This pattern of apparently overshooting carrying capacity following an initial release has been reported for a number of ungulates (Caughley, 1976). Disease may have caused the decline; predation and/or competition with wild horses (Equus caballus) may also have been factors. A spatial model of wild horse carrying capacity (Coughenour, 1999) was developed to assist managers in evaluating wild horse population numbers. Studies of summer consumption rates by wild horses and other ungulates were conducted by James Detling and students from Colorado State University (Peterson and others, 1997; Gerhardt and Detling, 1998; Fahnestock, 1998). Diets and habitat use of wild horses, bighorn sheep and mule deer were studied by Coates and Schemnitz (1989) and Kissell and others (1996). Both Coates and Schemnitz (1989) and Kissell and others (1996) reported a high degree of dietary overlap between wild horses and bighorn sheep. However, Kissell and others (1996) concluded that a high degree of spatial separation between wild horses and bighorn sheep, at least under the study conditions, precluded any significant negative competitive influences. The most important overlap of bighorn sheep and wild horses seems to be on winter range. Competition can be difficult to determine, since current conditions may not reveal competition that has already occurred. Managers remained concerned about the declining bighorn population for the following reasons: Our objectives for the study were to: Evaluate current disease problems and mortality rates, including predation rates, in bighorn sheep through captures and monitoring of bighorn sheep. Determine winter spatial relations of wild horses and bighorn sheep through aerial surveys of winter range. Conduct habitat measurements to determine why bighorn sheep were not using what had been mapped with a GIS-based habitat model as suitable bighorn sheep habitat, and sample winter and summer consumption rates of key forage species to see if there were other factors precluding use of these areas by bighorn sheep. Compare winter lamb recruitment rates and total vegetation consumption rates on bighorn sheep winter use areas, with and without wild horses present. Complete year-round GIS-based distribution maps for wild horses and bighorn sheep in order to identify key areas of species overlap and key areas of species separation for potential habitat improvements. NPS park staff asked USGS to answer the following questions: (a) how large is the population?; (b) how many rams are in the population?; (c) in addition to low lamb recruitment, is mortality or dispersal of adults likely contributing to the decline?; and (d) is trophy hunting of rams contributing to the decline?

Report

Restoration of bighorn sheep metapopulations in and near 15 national parks: Conservation of a severely fragmented species; Volume I, Planning, problem definition, findings, and restoration

Bighorn sheep ( Ovis canadensis ) were historically a ubiquitous species. Prior to the arrival of Europeans, they were seemingly widespread in nearly all steep habitats in the mountains, foothills, river breaks, and prairie badlands of the western United States. However, since catastrophic declines in the late 1800s and early 1900s, most extant populations have existed as small, isolated groups in a highly fragmented distribution. Stochastic events such as seasonal weather change or population fluctuations render these small populations more prone to extirpation than larger populations. Three different subspecies of bighorn sheep were eliminated from 14 of 18 National Park System (NPS) units in the 6-state Intermountain Region of the western United States (Singer 1994). In 1990, when this restoration was initiated, only 4 (18%) of 22 discrete park populations or metapopulations were considered large enough (300-500 animals) to be secure for long-term management. Five (23%) other populations numbered 100-299 animals and 3 (14%) populations numbered 75-99 animals. But, the remaining 10 populations (45%) were either extirpated (n = 2), remnant populations (n = 5, populations of 7-10 animals), or vulnerable to extirpation (n = 3, populations of less than 50 animals). Restoration prior to 1991-96 was largely completed in one NPS unit, but was incomplete in the remaining units. Most bighorn sheep are not federally listed as endangered or threatened species, although the California peninsular population of desert bighorn sheep was recently listed as endangered. The bighorn sheep is a rare or uncommon species that is declining in many parts of its range but is abundant in other areas and still relatively easy to study and manage. The Secretary of the Department of the Interior, Bruce Babbitt, directed the Biological Resources Division of the U.S. Geological Survey in 1993 (when this agency was still the National Biological Service) to research and recover species that were declining to avoid expensive and controversial federal listing. Because the capture and moving of the species are still relatively uncomplicated and because some source stocks are available, aggressive restoration in 15 National Park System units in the former Rocky Mountain Region was recommended in 1990. This report details the 7-year restoration of bighorn sheep to all currently suitable historic habitats in the national parks of the former Rocky Mountain Region (now the Intermountain and Midwest regions of the National Park Service). The purpose of the first phase of the restoration during 1991-93 was to conduct research and population surveys and to formulate the restoration plans. The purpose of the second phase of the initiative during 1994-97 was to conduct GIS-based habitat and biological assessments of prospective restoration sites, write restoration plans, and restore and monitor the released bighorn sheep.

Colorado, Montana, North Dakota, South Dakota, Uta

Restoration of bighorn sheep metapopulations in and near 15 national parks: Conservation of severely fragmented species; Volume II, Synopsis of research findings

In 1991, the National Park Service (NPS) initiated a series of research studies to support an effort to restore bighorn sheep (Ovis canadensis) into 15 national parks in the Intennountain West and Colorado Plateau areas (Fig. 1). The Biological Resources Division of the U.S. Geological Survey provided scientific advice and research coordination for the NPS restoration (Fig. 2). The research studies were conducted by scientists from the Biological Resources Division of the USGS (fonnerly NBS) (11 research studies), university-based scientists (Univ. of Wyoming- 2 studies, University of Colorado- 1, Colorado State University- 2, University of California, White Mountain Center- 1, Northern Arizona University - 1, Montana State University - 1) and by state agency veterinarians: Drs. Beth Williams of Wyoming, Mike Miller of Colorado, and Terry Spraker of Colorado State University. Only the highlights of these research studies are presented below. Full research reports are available in Volume III of this series.

Colorado, Montana, North Dakota, South Dakota, Uta

Caribou calf mortality in Denali National Park, Alaska

Calf mortality is a major component of caribou (Rangifer tarandus) population dynamics, but little is known about the timing or causes of calf losses, or of characteristics that predispose calves to mortality. During 1984-87, we radiocollared 226 calves (≤3 days old) in the Denali Caribou Herd (DCH), an unhunted population utilized by a natural complement of predators, to determine the extent, timing, and causes of calf mortality and to evaluate influences of year, sex, birthdate, and birth mass on those losses. Overall, 39% of radio-collared calves died as neonates (≤15 days old), and 98% of those deaths were attributed to predation. Most neonatal deaths (85%) occurred within 8 days of birth. Few deaths occurred after the neonatal period (5, 10, and 0% of calves instrumented died during 16-30, 31-150, and >150 days of age, respectively). Survival of neonates was lower (P = 0.038) in 1985, following a severe winter, than during the other 3 years. In years other than 1985, calves born during the peak of calving (approx 50% of the total, born 5-8 days after calving onset) experienced higher (P < 0.001) neonatal survival than did other calves. Grizzly bears (Ursus arctos), wolves (Canis lupus), and unknown large predators (i.e., grizzly bears or wolves) accounted for 49, 29, and 16% of the neonatal deaths, respectively. The rate of bear-caused mortalities declined (P < 0.001) with calf age, and bears killed few calves >10 days old. Wolf predation was not related (P > 0.05) to calf age and peaked 10 days after onset of calving. Grizzly bear and wolf predation on neonates during the calving season was a limiting factor for the Denali Caribou Herd.

Journal of Wildlife Management

Niche relationships within a guild of ungulate species in Yellowstone National Park, Wyoming, following release from artificial controls

Niche relationships and diet overlaps were compared among elk (Cervus elaphus), bison (Bison bison), bighorn sheep (Ovis canadensis), mule deer (Odocoileus hemionus), and pronghorn antelope (Antilocapra americana) between 1967–1970 and 1986–1988, a period when total ungulate numbers nearly tripled on Yellowstone's northern range. Ungulate species ratios on Yellowstone's northern winter range during the latter period were 100 elk : 10 mule deer : 3 bison : 2 pronghorns : 1 bighorns. Elk numbers were positively correlated to bison, mule deer, and pronghorn numbers (r2 = 0.76, 0.97, and 0.48, respectively, P < 0.01). Few other changes in habitat use or habitat overlap occurred, and diets for only 2 of the 10 species pairs, elk-bighorn (Spearman's rank order coefficient (RHO) = 0.55, P < 0.05) and mule deer – pronghorn (RHO = 0.64, P < 0.05), were significantly associated with each other. Bison consumed more grass and fewer sedges, mule deer more fringed sage (Artemisia frigida) and more rabbit-brush (Chrysothamnus spp.), and bighorn sheep more grasses and fewer sedges, while pronghorns ate less saltsage (Atriplex nuttalli) but more big sagebrush (Artemisia tridentata) during 1986–1988 than during 1967–1970. Bison expanded their range and bison and bighorn sheep used a wider variety of habitats. We found little evidence of change in competitive interactions between ungulate species. A few diet and habitat overlaps increased, the opposite of the prediction from the competitive exclusion principle amongst species, suggesting that intraspecific competition was more important. Several explanations are proposed for the lack of changes in niche relationships during a period of near tripling in density of the ungulate guild.

Wyoming

Physiological responses of Yellowstone bison to winter nutritional deprivation

Because nutrition is critically related to other aspects of bison ( Bison bison ) ecology, and the winter ranges inhabited by bison in Yellowstone National Park (YNP) are ecologically diverse, it was important to determine if nutritional deprivation differences occurred among winter ranges. We used chemistry profiles of urine suspended in snow to compare nutritional deprivation of bison from January to April 1988 on 4 sampling areas of 3 winter ranges in YNP. Declining (P < 0.001) trends of urinary potassium: creatinine ratios in bison on all 4 sampling areas indicated progressive nutritional deprivation through late March. Concurrent increases (P ≤ 0.001) in mean urea nitrogen: creatinine ratios from late February through late march in 3 of 4 areas suggested that increased net catabolism was occurring. Diminished creatinine ratios of sodium and phosphorus reflected low dietary intake of these minerals throughout winter. Mean values and trends of urinary characteristics indicated nutritional deprivation varied among 3 winter ranges in YNP. Continued physiological monitoring of nutritional deprivation, along with detailed examination of other aspects of the bison's ecology, will provide greater insight into the role of ungulate nutrition in the dynamics of such a complex system and improve management.

Journal of Wildlife Management

Potential ungulate prey for Gray Wolves

Data were gathered for six ungulate species that reside in or near Yellowstone National Park. If gray wolves ( Canis lupus ) are reintroduced into the Yellowstone area, their avoidance of human activities or their management by human may determine their range. Therefore, the area of wolf occupation cannot be predicted now. We restricted our analysis to Yellowstone National Park and to the adjacent national forest wilderness areas. We included mostly ungulate herds that summer inside or adjacent to the park and that would probably be affected by wolves. Our wolf study area includes Yellowstone National Park and adjacent wilderness areas most likely to be occupied by wolves. We reviewed publications, park records, survey reports, and state fish and game surveys and reports for statistics on ungulate populations. These data [provide an overview of ungulate populations and harvests. Each ungulate herd is described in detail. We restricted our analysis to 1980-89, because population surveys were more complete during that period and because population estimates of most ungulate populations had increased by the 1980's. We feel the higher estimates of the 1980's reflect more up-to-date techniques and are most representative of the situation into which the wolves would be reintroduced.

Idaho, Montana, Wyoming

Using Pop-II models to predict effects of wolf predation and hunter harvests on elk, mule deer, and moose on the northern range

The effects of establishing a gray wolf ( Canis lupus ) population in Yellowstone National Park were predicted for three ungulate species—elk ( Cervus elaphus ), mule deer ( Odocoileus hemionus ), and moose ( Alces alces )—using previously developed POP-II population models. We developed models for 78 and 100 wolves. For each wolf population, we ran scenarios using wolf predation rates of 9, 12, and 15 ungulates/wolf/year. With 78 wolves and the antlerless elk harvest reduced 27%, our modeled elk population estimated were 5-18% smaller than the model estimate without wolves. With 100 wolves and the antlerless elk harvest reduced 27%, our elk population estimated were 11-30% smaller than the population estimates without wolves. Wolf predation effects were greater on the modeled mule deer population than on elk. With 78 wolves and no antlerless deer harvest, we predicted the mule deer population could be 13-44% larger than without wolves. With 100 wolves and no antlerless deer harvest, the mule deer population was 0-36% larger than without wolves. After wolf recovery, our POP-II models suggested moose harvests would have to be reduced at least 50% to maintain moose numbers at the levels predicted when wolves were not present. Mule deer and moose population data are limited, and these wolf predation effects may be overestimated if population sizes or male-female ratios were underestimated in our population models. We recommend additional mule deer and moose population data be obtained.

Idaho, Montana, Wyoming