Browsing by Author "Vaughan, Michael R."

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    Aspects of Reproduction and Cub Survival in a Hunted Population of Virginia Black Bears
    Echols, Kim Needham (Virginia Tech, 2000-06-09)
    We measured black bear (Ursus americanus) reproduction and cub survival during 1994 - 1998, and 1995 - 1999, respectively, in the George Washington and Jefferson National Forests in Virginia to determine age-specific and overall cub production and cub survival. We observed females in estrus between 6 June and 22 August; the mean date of estrus was 17 July. Ages of primiparity ranged between 3 and 5 years with an average of 3.36 years (n=11, SE=0.15). Average litter size for 1995 - 1998 was 2.32 cubs/litter (SE=0.11, n=53) and 85.7% of available females ≥ age 4 (those not accompanied by cubs) reproduced in a given den season. We monitored 98 (48M:50F) black bear cubs equipped with expandable radio-collars (Higgins 1997) or radio transmitters implanted subcutaneously between 1995 and 1999 to estimate cub survival. Kaplan-Meier staggered entry analysis provided 306-day survival rates for 82 cubs. The survival estimates for males and females were 73% (0.49, 0.96) and 91% (0.80, 1.00), respectively. The overall 306-day survival rate for all cubs was 81% (0.67,0.94) using Kaplan-Meier and 76% (0.63, 0.92) using Heisey-Fuller (Mayfield) methods. We also evaluated the utility of radio transmitters implanted subcutaneously in 42 (21M:21F) wild black bear (Ursus americanus) cubs from 2 study areas in Virginia between 1996 and 1999 to monitor first year cub survival. More than 64% (27 of 42) of the implants fell out prematurely (2-198 days), and 16.6% (7 of 42) failed for unknown reasons. Less than 5% (2 of 42) of these cubs denned wearing failed implants, and 9.5% (4 of 42) experienced mortality less than 1 month after implant surgery. About 9.5% (4 of 42) of implanted black bear cubs wore working transmitters through to the following den season.
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    An assessment of Quality Deer Management on a private hunt club in the Virginia Piedmont
    Batts, Gregory K. (Virginia Tech, 2008-02-15)
    I examined the efficacy of Quality Deer Management (QDM) on Amelia Springs hunt club in Amelia County, Virginia, during 2003-2006. I examined home range dynamics of male white-tailed deer (Odocoileus virginianus), deer/hunter interactions, and aspects of population dynamics. I also developed a new rocket net method to capture deer using a remote video system that was more efficient than traditional methods. I monitored 20 deer; 50% died due to hunting and 15% to natural mortality. The emigration rate for juvenile males was 46%, dispersal distance averaged 6.4 km. I used Home Range Extension (HRE) in ArcView to generate annual home ranges (adaptive-kernel) for 16 male deer; I also generated annual and seasonal home ranges using MCP. Annual and seasonal home ranges (MCP) of adult males were larger than those of juveniles. Adult male annual home ranges averaged 2.5 km2 and juveniles 0.9 km2. Seasonal home ranges of adult males were 1.6 km2 and 1.3 km2 during non-hunting and hunting seasons respectively. Juvenile non-hunting and hunting season home ranges were 0.6 km2 and 0.8 km2 respectively. I detected no differences in day/night movements of male deer during the hunting season; however, deer appeared to avoid areas that were hunted based on hunter GPS locations and deer locations during the hunting season. Frequency of deer movement increased during October-November. Population estimates based on remote camera mark-recapture averaged 60 antlered males for the 3-year survey period. Using population reconstruction, the minimum buck:doe ratio was 1:1.8. Estimated density of antlered males was 4.1/km2, in Amelia County, and 5.0/km2 for Amelia Springs. Deer harvested on Amelia Springs, compared to deer harvested on other hunt clubs in Amelia County, were larger. Antler diameters averaged 32.6mm on Amelia Springs versus 26.9mm for other Amelia county hunt clubs, average age at harvest for 2+ males was higher on Amelia Springs (2.4) than other Amelia county hunt clubs (2.2), and dressed body weights averaged 11.2kg heavier (46.2 kg versus 35 kg) on Amelia Springs. QDM on Amelia Springs appears to be successful based on the results. While bigger bucks existed on Amelia Springs, hunters failed to encounter them. Hunters likely would increase buck sightings during the hunting season by becoming more mobile. Expectations of the size of animal (antlers) Amelia Springs can produce should be adjusted to reflect what is possible based on the habitat. The harvest program in place should be continued at the current level for continued success using QDM.
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    Black bears in Canyon de Chelly National Monument: Life in a changing environment
    Tredick, Catherine Anne (Virginia Tech, 2011-06-23)
    Understanding how wildlife utilize habitat at varying scales is important for understanding and predicting potential impacts of landscape changes (e.g., habitat loss and fragmentation, restoration efforts, climate change, etc.) and in determining effective strategies for conservation and management. This research examines fine-scale and landscape-level habitat use of black bears in Canyon de Chelly National Monument (CACH), Arizona, USA in the context of large-scale landscape change. Currently, CACH is undergoing a large-scale restoration effort to remove all of the non-native Russian olive (Elaeagnus angustifolia) and tamarisk (Tamarix ramosissima and T. chinensis) within the monument. As black bears rely on the Russian olive as a primary food source, a major goal of this research was to evaluate how bears currently use this resource and how its removal might affect bear habitat use and bear-human interactions within the monument. I developed a model of 3rd order (fine-scale) black bear habitat use in CACH using an occupancy modeling approach. Model results indicated that fine-scale habitat selection by bears in CACH is being driven by the presence of non-native Russian olive as well as native food sources. Thus availability of native foods may be sufficient to maintain the bear population in CACH, and bears may quickly adapt to the loss of the non-native food source. Similarly, results showed that bears avoid human areas and farmsteads and prefer higher elevations, suggesting that once olive is removed in the lower canyons near human areas, bear-human interactions may become less of a problem over the long term. I also developed a model of 2nd order (landscape-level) habitat use and evaluated movement patterns of black bears in CACH using location data collected from GPS collars. Model results showed that bears selected areas with higher tree canopy cover and terrain ruggedness, indicating that forest cover and escape cover are primary factors driving black bear habitat selection at the landscape scale in this region. Movement patterns revealed large mean daily movements and low average turning angles, indicating long, linear movements designed to take advantage of the mosaic of available habitats and food resources available over larger areas. I extracted DNA from hair samples collected throughout the study area to examine genetic variability and population structure of black bears in the region. Analyses revealed a relatively healthy, panmictic population across the wider landscape. No substantial genetic structuring was observed in multiple analyses, though I did find evidence of a slight isolation-by-distance pattern within the population. Measures of both current (Nb = 24) and long-term (Ne = 579) effective population size indicated a relatively high number of breeders in the current population and a sufficient amount of gene flow within the larger "superpopulation" to maintain long-term genetic viability. I focused the final portion of my dissertation research on understanding the factors that influence stakeholder acceptance capacity for black bears, which plays a central role in contemporary wildlife management issues, including human-wildlife conflict. I used data from telephone interviews of 1,546 residents in Virginia to develop conceptual models of black bear acceptance capacity at both county and state levels. Model results suggested that more deep-seated and less easily influenced factors (e.g., values and risk perceptions) are at the heart of stakeholder attitudes and perceptions of wildlife, making influencing these perceptions more difficult. Agencies can indirectly affect these attitudes and perceptions, however, by targeting more potentially pliable factors related to wildlife acceptance capacity such as knowledge, personal experience with wildlife, and trust in management agencies.
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    Contaminants at a Shooting Range: Toxicological and Nutritional Significance to Birds and Mammals
    Gonzalez, Gabriela Rae (Virginia Tech, 2003-08-08)
    Target shooting in the United States has become an increasingly popular sport in the last century. In addition to the large quantity of lead pellets littering range grounds and surrounding land, considerable amounts of clay target fragments cover shooting range areas as well as adjoining habitats. Polycyclic aromatic hydrocarbons (PAHs) within the target, as well as lead pellets from shotguns pose multiple threats to a variety of wildlife. To determine the effects of clay target and lead pellet ingestion on wildlife, I conducted controlled experiments on Coturnix quail exposed to clay targets in the lab, and collected wild birds and mammals exposed to lead pellets at a shooting range. The first Coturnix study determined whether quail voluntarily consumed target fragments or limestone fragments. In both fall (F=29.2, P<0.01) and spring (F=6.45, P=0.02) experiments, I found that quail consistently selected limestone fragments, but almost completely rejected clay target fragments. In the second study, quail were force-fed varying amounts of target dust on a weekly basis to simulate sporadic exposure to clay target dust. In both summer (F=1.63, P=0.23) and winter (F=0.34, P=0.8) trials, male quail did not have significant weight loss. Female quail had insignificant weight losses in summer trials (F=1.63, P=0.23) but experienced weight gains in winter trials (F=3.53, P=0.04). In the third and final Coturnix study, varying amounts of target dust were incorporated into daily feed rations to simulate frequent exposure to clay target dust. Male quail experienced weight loss in both summer (F=16.13, P<0.01) and winter (F=8.47, P<0.01) trials. Female quail also suffered weight loss in both summer (F=15.62, P<0.01) and winter (F=17.50, P<0.01) trials. Weight loss likely resulted from inadequate nutrition as opposed to target poisoning. However, because there were no biochemical analyses performed to test for PAH presence, no definite conclusions can be made. The second study focused on lead contamination in Passeriformes, perching birds, and small mammals. Seventeen of 20 birds (85%) (Passerine spp) captured at the shooting range had elevated lead levels (F=5.21, P<0.028), when compared to birds (n=20) at the control site. Nine of 26 (35%) white-footed mice (Peromyscous leucopus), trapped at the shooting range had elevated liver (F=9.78, P=0.0029) and kidney (F=22.49, P<0.01) lead levels. These results indicate that Passerine species as well as Peromyscous species around shooting ranges inadvertently consume lead, either as lead pellets, mistaking them for grit or dietary items, or through environmental sources such as water, soil, and vegetation.
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    Den Tree Avaliability and Denning Success of Black Bears on Industrial Forest Lands and National Forest Lands in Virginia and West Virginia
    Quince, Gyasi Adrian (Virginia Tech, 2002-06-05)
    During 1999-2001, potential den trees and denning success of black bears was investigated on industrial forest lands and national forest lands in Virginia and West Virginia. One hundred and fifty seven potential den trees were found on five study areas, 135 on national forest lands and 22 on the industrial forest lands. Twenty-seven hollow potential den trees were found, 22 on national forest lands and 5 on industrial forest lands. Densities of potential den trees ranged from 12.5/km2 to 437.5/km2 on the national forest lands and on the industrial forest lands the densities ranged from 0-187.5/km2. There were significantly higher densities (F1, 69=5.86, P=0.0181) of potential den trees on the national forest land than on the industrial forest land. There were also significantly (F=2, 68=7.86 P=0.0008) higher densities of hollow potential den trees based on the stand age class. Denning success for females expected to have cubs was 98% (n=46) in tree dens on national forest lands and 100% (n=8) on industrial forest lands. Denning success for females expected to have cubs was 76% (n=55) in ground dens on national forest lands and 80% (n=5) in ground dens on industrial forest lands. Females expected to have cubs on the national forest lands had an overall denning success of 89% (n=80) and on industrial forest lands there was overall, 92% (n=13) denning success. Chi-square tests showed that abandonment of dens by bears was significantly higher (x2=19.02, 1df) in ground dens than in tree dens. Litter sizes were not different from national forest lands and industrial forest lands (t=-0.36, 44df, P=0.84). The mean litter sizes on the national forest lands was 2.55 (SE=0.16) cubs per litter and on the industrial forest land 2.4 (SE=0.22) cubs per litter.
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    Densities, habitat-use, and mesopredator release of the ocelot in Belize
    Davis, Miranda L. (Virginia Tech, 2008-05-04)
    The mesopredator release hypothesis suggests that small carnivore populations are negatively affected by competition with larger carnivore species; this could hinder the conservation of mesopredators in areas where large top-carnivores are prioritized. I investigated the ecology of ocelots in Belize and examined the role of mesopredator release in the neotropical carnivore guild to determine if ocelots are limited by competition with jaguars and pumas. I conducted remote camera surveys and sampled habitat within four protected areas: three broadleaf sites and one pine forest site. I measured ocelot activity and habitat-use with respect to the activity of jaguars and pumas at one broadleaf site and one pine forest site; additionally, I calculated estimates of ocelot and jaguar densities within those two sites. Ocelot presence was positively related to jaguar activity in the pine forest and to both jaguar and puma activity in the broadleaf forest. There were few relationships with habitat characteristics; however, in the broadleaf site, ocelot activity was positively related to road width and large avian prey activity. Both jaguar and ocelot densities were low in the pine forest and higher in the broadleaf site. Preliminary findings from the remaining two sites suggest that future results may be similar to those from the first two sites analyzed. I conclude that ocelot populations are not negatively affected by jaguars and that a negative effect of pumas is unlikely. Results imply that mesopredator release does not limit ocelot populations in these areas.
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    Digestibility of Foods and Anthropogenic Feeding of Black Bears in Virginia
    Gray, Rachel Masterson (Virginia Tech, 2001-08-10)
    For years, bear hunters have provided an unknown amount of food to black bears (Ursus americanus) in Virginia, supplementing their natural food supply. Possible effects of feeding bears can be negative, such as food conditioning and habituation to people, or positive, such as enhanced or sustained reproductive rates, especially in years of mast crop failure. In July 1999, the Virginia Department of Game and Inland Fisheries (VDGIF) adopted a regulation recommendation that prohibited the feeding of wildlife on national forest and department-owned lands. We mailed a survey to all members of the Virginia Bear Hunters Association (VBHA, n = 459) to determine the amount of food provided to bears by bear hunters in Virginia between 1 July 1998 and 30 June 1999. Survey response rate was 52%. One hundred thirteen of 238 (47.5%) survey respondents spent over $18,000 on supplemental food in one year, averaging $163/person. One hundred twenty-eight respondents reported cumulatively providing nearly 3 million kilograms of food to bears between 1 July 1998 and 30 June 1999. Whole-shelled corn, pastries, and grease accounted for 58% of the total food by weight; however, whole-shelled corn, pastries, and bread were the 3 most common foods offered. Feeding occurred in 25 counties in Virginia, primarily during July, August, and September. Rockingham County had the highest proportion of feeders (47%), followed by Giles (15%), Augusta (14%), Craig (9%) and Montgomery (9%) Counties. Accordingly, Rockingham County received the highest proportion of food (42%), followed by Craig (8%), Giles (7%), Montgomery (6%), and Tazewell (6%) Counties. The low survey response rate, coupled with the fact that 48% of sampled nonrespondents fed bears, suggests that the total amount of annual feeding in Virginia may exceed the total determined during this study. However, the possibility of an avidity bias, in which a higher response by the more avid feeders would erroneously inflate a total estimate extrapolated to include feeding by nonrespondents, cannot be eliminated. Little work has been done pertaining to digestibilities of black bear foods, particularly those in the eastern United States. We determined digestibilities for several important eastern black bear foods, including northern red oak (Quercus rubra), white oak (Q. alba), and chestnut oak acorns (Q. prinus), squawroot (Conopholis americana), high-protein dogfood, shelled corn, and doughnuts. All diets were evaluated for nutrient content as well as percent dry matter digestibility of crude protein, ether extract (a measure of fat), and gross energy. Diets we suspected were high in fiber were additionally evaluated for neutral detergent fiber or acid detergent fiber content and digestibility. Feeding trials were conducted with 1 male and 4 female captive bears during fall 1998, and 5 female captive bears during fall 1999, at the Center for Ursid Research, Blacksburg, Virginia. Acorns and squawroot were high in fiber (45-62%) and moderate to low in protein (5-7%). Fat content was high in doughnuts (24.7) and northern red oak acorns (12%), moderate in dogfood (8.2%) and white oak acorns (4.7%), and relatively low in shelled corn, chestnut oak acorns, and squawroot (1.3-3.5%). Crude protein was high in dogfood (29%) and moderate in all other diets (4.7-8.8%). Neutral detergent fiber content was high in squawroot and all acorn types (45-62.4%). Acid detergent fiber was high in squawroot (47%) and moderate in dogfood (9%). In 1998, crude protein digestibility was high in doughnuts (86%), moderate in shelled corn and white oak acorns (50-64%), and negligible in northern red oak acorns (-39%). Ether extract digestibility was high in doughnuts, red oak acorns, and white oak acorns (80-97%), and moderate in shelled corn (67%). Fiber digestibility was tested only on northern red oak and white oak acorns, and was moderate (62% and 68%, respectively). In 1999, crude protein digestibility was high in dogfood, dogfood mixed with squawroot, and doughnuts (75-85%); moderate in shelled corn, squawroot, and white oak acorns (43-58%); and low to negligible in chestnut oak and northern red oak acorns (-13-6%). Ether extract digestibility was moderate in shelled corn (64%) and squawroot (60%), and high in all other diets (79-97%). Neutral detergent fiber digestibility was moderately high in all acorn types (54-71%). Acid detergent fiber digestibility was moderate for dogfood (66%), and associative effects occurred between the fiber in squawroot and dogfood, resulting in negative fiber digestibility in squawroot determined by difference (-19%). A luxury not always realized in wildlife management is the ability to investigate possible effects of management or policy changes prior to their employment. The amount of food provided to bears by bear hunters in Virginia may have been more substantial than previously believed, and likely provided bears with a high energy, stable food source that supplemented their natural food supply. We did not have the opportunity to study the effects of removing that food source on public lands prior to the regulation change; however, the opportunistic feeding strategy of bears, coupled with sufficient mast production at the time of the regulation change, likely softened any negative effects that may have occurred as a result of the removal of supplemental food as a regular food source for some bears.
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    Distribution and Relative Abundance of Nutrient Transporter mRNA in the Gastrointestinal Tract of Black Bears
    Gilbert, Elizabeth R. (Virginia Tech, 2005-08-08)
    Black bears are omnivorous, and tend to be opportunistic feeders, in that they will eat what is readily abundant or available. The end-products of intestinal digestion are absorbed by the body through the action of transporter proteins expressed on the brushborder membrane of small intestinal epithelial cells. The goal of this study was to increase the understanding of the physiological processes associated with nutrient assimilation by black bears. Distribution and relative abundance of mRNA of a peptide transporter (PepT1), a glucose transporter (SGLT1), two AA transporters (NBAT, bo,+AT), and a digestive enzyme, aminopeptidase N (APN), in the intestinal tract of black bears were investigated. Ten bears were used for this study. For tissue collection, the intestine was removed from the animal and divided into five sections. Each collected section was opened longitudinally, rinsed in ice-cold PBS, and the mucosal scrapings were stored at -80C. Total RNA was extracted and quantified by spectrophotometry. Abundance of PepT1, SGLT1, NBAT, bo,+AT, and APN mRNA was determined by performing Northern blots, using bear cDNA probes. Northern blot data were quantified by densitometric analysis, with the abundance of each gene expressed relative to GAPDH. Abundance of PepT1 (P < 0.05), APN (P < 0.05), and SGLT1 (P < 0.0001) changed quadratically from the proximal to the distal intestine with abundance being greatest in the midregion. Abundance of bo,+AT mRNA increased linearly (P < 0.05) from the proximal to distal intestine. Abundance of NBAT mRNA did not change among intestinal segments.The absolute number of molecules of mRNA/ng of total RNA for each gene was determined using Real-Time PCR. Similar to the Northern results, abundance of PepT1 (P < 0.0003), SGLT1 (P < 0.0003), and APN (P < 0.02) changed quadratically from the proximal to distal intestine with abundance being greatest in the mid-region, and bo,+AT mRNA increased linearly (P < 0.0001) from the proximal to distal intestine. NBAT mRNA abundance also increased linearly (P < 0.0001) from proximal to distal intestine. PepT1 mRNA was present at tenfold or greater levels than AA transporter mRNA in all segments of the intestine, suggesting that di- and tripeptides constitute the major form in which AAs are absorbed. NBAT and bo,+AT mRNA abundance was greater towards the distal portion of the intestine, suggesting their importance in salvaging remaining unabsorbed AAs.These results indicate that the mRNA of nutrient transporters examined and APN are differentially expressed throughout the gastrointestinal tract of black bears, suggesting their involvement in nutrient assimilation.
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    DNA-based hair sampling to identify road crossings and estimate population size of black bears in Great Dismal Swamp National Wildlife Refuge, Virginia
    Wills, Johnny (Virginia Tech, 2008-08-29)
    The planned widening of U.S. Highway 17 along the east boundary of Great Dismal Swamp National Wildlife Refuge (GDSNWR) and a lack of knowledge about the refuge's bear population created the need to identify potential sites for wildlife crossings and estimate the size of the refuge's bear population. I collected black bear hair in order to collect DNA samples to estimate population size, density, and sex ratio, and determine road crossing locations for black bears (Ursus americanus) in GDSNWR in southeastern Virginia and northeastern North Carolina. I also investigated bear/vehicle collisions to determine patterns of road crossing. Genetic analysis of 344 hair samples collected on 2 trapping grids identified 85 unique individuals which I used in a mark-recapture analysis. Estimated population size on the trapping grids was 105 bears (95% CI = 91-148) and average density was 0.56 bears/km². This density estimate projected over the entire Great Dismal Swamp ecosystem yielded a population estimate of 308 bears (550 km2 X 0.56 bears/km²). Similar population estimates generated by Hellgren (1988), Tredick (2005), and this study suggest a stable bear population in the Great Dismal Swamp ecosystem over a 20-year period. I erected a 2.3-kilometer long strand of barbed wire along U. S. Highway 17 to monitor road crossing patterns near the Northwest River drainage. Genetic analysis identified 6 bears (4 males, 1 female, 1 unknown) that apparently crossed the highway in a 10-month period. Five of 6 bears deposited hair in a 171-m section which included the Northwest River corridor. The 6 bears detected crossed the road at least 11 times. I investigated 10 reports of bear/vehicle collisions on the periphery of the refuge from June 2000 to May 2002. Six bears (4M:1F:1 unknown) were confirmed killed during this time period. Based on reported bear/vehicle collisions from Hellgren (1988), the Virginia Department of Game and Inland Fisheries database, and this study, a minimum of 4 to 5 bears are struck by vehicles each year on the periphery of the refuge. I identified 2 areas of multiple bear/vehicle collisions: highway 58 on the north side of the refuge near Hampton Airport and Highway 17 on the eastern side of the refuge in the vicinity of the Northwest River corridor.
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    Ecology and conservation of Formosan clouded leopard, its prey, and other sympatric carnivores in southern Taiwan
    Chiang, Po-Jen (Virginia Tech, 2007-11-14)
    During 2000-2004 I studied the population status of the Formosan clouded leopard (Neofelis nebulosa brachyurus) and the ecology of its prey and other sympatric carnivores in the largest remaining lowland primary forest in southern Taiwan. My research team and I set up 232 hair snare stations and 377 camera trap sites at altitudes of 150-3,092m in the study area. No clouded leopards were photographed in total 13,354 camera trap days. Hair snares did not trap clouded leopard hairs, either. Assessment of the prey base and available habitat indicated that prey depletion and habitat loss, plus historical pelt trade, were likely the major causes of extinction of clouded leopards in Taiwan. Using zero-inflated count models to analyze distribution and occurrence patterns of Formosan macaques (Macaca cyclopis) and 4 ungulates, we found habitat segregation among these 5 herbivore species. Formosan macaques, Reeve's muntjacs (Muntiacus reevesi micrurus), and Formosan serows (Nemorhaedus swinhoei) likely were the most important prey species of Formosan clouded leopards given their body size and high occurrence rates in lower altitudes. In contrast, sambar deer (Cervus unicolor swinhoii) tended to occur more frequently as altitude increased. Formosan macaques exhibited seasonal differences in occurrence rates and were absent at altitudes > 2,500m in winter. Only Formosan serows showed preference for cliffs and rugged terrain, while the other 4 species, except wild boars (Sus scrofa taivanus), avoided these areas. Habitat segregation in forest understory and structure were more pronounced among the 4 ungulates. Forest structure rarely affected occurrence rates of Formosan macaques on the ground. Niche relationships of the other sympatric carnivores were studied through habitat, diet, and temporal dimensions. Resource partitioning by carnivores was observed. Altitude was the strongest factor explaining the composition of the carnivore community in the local study-area scale and in the landscape scale across Taiwan. Carnivores could be divided into 2 groups: low-mid altitude consisting of Formosan ferret badgers (Melogale moschata subaurantiaca), gem-faced palm civets (Paguma larvata taivana), lesser oriental civets (Viverricula indica taivana), crab-eating mongooses (Herpestes urva formosanus), leopard cats (Prionailurus bengalensis chinensis), and feral cats (Felis catus), and the mid-high altitude group consisting of yellow-throated martens (Martes flavigula chrysospila), Siberian weasels (Mustela sibirica taivana), and Asiatic black bears (Ursus thibetanus formosanus). Carnivore richness was higher at mid altitudes where these 2 groups overlapped (i.e. mid-domain effect). The low-mid altitude carnivores were more nocturnal and tolerant of human activity and forest alteration except crab-eating mongooses, which were diurnal and avoided human encroachment. Similar to crab-eating mongooses, the mid-high altitude carnivores also avoided human encroachment and were diurnal except for Siberian weasels, which were more nocturnal. Diet summary based on their major food items for all sympatric carnivores revealed 3 groups of foragers which foraged on: invertebrates, small mammals, and plant fruits. Felidae, yellow-throated martens, and Siberian weasels preyed on small mammals. Asiatic black bears and gem-faced palm civets ate mostly plant fruits. The other 3 carnivores were mainly invertebrate foragers. These 9 carnivores partitioned resource uses in the 3 niche dimensions except for some overlap in resource use by leopard cats and feral cats. Prey base for Formosan clouded leopards and the carnivore richness in Taiwan were found to be lower in areas with higher levels of human activity. On the other hand, Formosan macaques and ungulates could become over-abundant without human hunting and top carnivore predation. Mesopredator release may occur because of vanishing top carnivores, causing reduction of the lower trophic level prey species. It is important to assess the cascading impacts of the loss of the Formosan clouded leopards and Eurasian otters (Lutra lutra chinensis) and the declining Asiatic black bears and to consider reintroduction of Formosan clouded leopards, as well as active management of the other larger mammals. These results provided baseline information for reintroduction of clouded leopards and management of their prey and generated new hypotheses regarding the ecology of these large mammals for future investigation.
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    Ecology and restoration of Sumatran tigers in forest and plantation landscapes
    Sunarto, Sunarto (Virginia Tech, 2011-02-11)
    Tigers (Panthera tigris Linnaeus, 1758) are in danger of extinction. Their populations have declined from ~100,000 to only ~3,000 individuals in a century and their habitat has shrunk to less than 7% of the historic range. Of the five extant tiger subspecies, the Sumatran tiger (Panthera tigris sumatrae Pocock, 1929) is the most seriously threatened. Currently determined as Critically Endangered under IUCN criteria, the Sumatran tiger is likely to become extinct unless effective conservation measures are enacted. Threats to the tiger include habitat destruction, killing due to conflict with humans and livestock, and poaching for illegal wildlife trade. Long-term survival of Sumatran tigers depends largely on the effectiveness of current conservation efforts in every tiger landscape. Successful conservation and management require accurate information on ecology of the species upon which decisions can be based. This study investigated basic ecological aspects of tigers and developed strategies for management and restoration to improve tiger viability in the Central Sumatra landscape. This landscape is comprised of natural forests and plantations managed for timber and agricultural commodities. The first chapter assesses the variation in tiger abundance across forest types in Southern Riau, and over time in Tesso Nilo National Park, all in Central Sumatra. Using camera traps, my team and I systematically sampled five blocks representing three major forest types in the region: peat land, flat lowland, and hilly lowland. I found that tiger abundance varied by forest type and through time. Excluding two sampling blocks where no tigers were photographed, the lowest tiger density was in peat land forest of Kerumutan, and the highest density was in the flat lowland forest of Tesso Nilo. Repeated sampling in the newly established Tesso Nilo National Park documented a trend of increasing tiger density (SE) from 0.90 (0.38) individuals/100 km2 in 2005 to 1.70 (0.66) individuals/100 km2 in 2008. Overall, tiger densities from this study were lower than most previous estimates from other parts of Sumatra. The trend of increasing tiger density in Tesso Nilo, however, suggests that the tiger population could be augmented by protection of habitats that were previously logged and severely disturbed. The second chapter examines the occupancy and habitat-use of the tiger across the major landcover types (natural forest, acacia plantation, oilpalm plantation, rubber plantation, and mixed agriculture). I found that tigers used some plantation areas, although they significantly preferred forests over plantations. In all landcover types, sites with tiger detections had thicker understory cover than sites without tiger detection. Modeling tiger occupancy while recognizing that probability of detection is not always perfect, I found that tiger occupancy covaried positively and significantly with altitude and negatively, but not significantly, with distance-to-forest-cores. Probability of habitat use by tigers covaried positively and significantly with understory cover and altitude, and negatively and significantly with human settlement and landcover rank. The results suggested that with adjustments in plantation management, tigers could use or roam through plantations within the habitat mosaic provided that the plantations had adequate understory cover and low level of human activity. They also could use riparian forests (as corridors) and smaller forest patches (as stepping stones) to travel between the main habitat patches across the forest and plantation landscape. The third chapter investigates the ecological characteristics and possible inter-specific interactions among wild felids, including tigers and smaller cats, based on data collected using systematic camera trapping in combination with information on their natural history. I found that despite overlap in resource needs of the five felid species, each appears adapted to specific environmental conditions allowing coexistence with other felids. The five felid species used statistically different elevations, with the golden cat found to inhabit the highest elevation. Two-species occupancy models showed that only leopard cats were found to co-occur with other felid species more frequently than expected by chance under independence. Species of similar size or eating similar-sized prey generally tended to have low coefficients of temporal activity overlap, suggesting avoidance. Temporal avoidance is likely occurring in three pairs of felids, namely clouded leopards and golden cats, clouded leopards and marbled cats, and marbled cats and leopard cats. Based on the differences in morphological and ecological characteristics, and on patterns of spatial and temporal occurrence, I identified six possible mechanisms by which felids in Central Sumatra maintain coexistence. I discussed the implications of this study for management, focusing on how to balance diversity and abundance of felids. The fourth chapter presents the tiger distribution models as a case study to illustrate the importance of accounting for uncertainty in species distribution mapping. I applied four modeling approaches, differing in how the response variable (tiger presence) is constructed and used in the models. I compared the performance and output of different models based on the relative importance of variables, descriptive statistics of the predictions, cross comparison between models using an error matrix, and validation using tiger presence data collected from independent surveys. All models consistently identified forest area within the grid as one of the most important variables explaining tiger probability of occurrence. Three models identified altitude as another important factor. While the four models were consistent in predicting relatively high probability of tiger occurrence for high elevation forest areas such as Rimbang Baling and Bukit Tigapuluh, they generally had a lower level of agreement in predictions for low elevation areas, particularly the peat land in the northeastern part of the study area. Based on the results of cross evaluation of the predictions among models and validation with the independent data, I considered the occupancy model to be superior to the others. If data collection format permits, I advocate the use of occupancy instead of the other modeling techniques to develop predictive species distribution maps. The last chapter constructs a strategy to restore the tiger population across the ecosystem of Central Sumatra through integration of knowledge on tiger ecology from previous chapters with consideration of the ecological conditions of the landscape in the region. The strategy combines existing knowledge of tiger conservation and regional ecosystem restoration. It recognizes the limitations and challenges of traditional nature protection and considers existing and new opportunities. Emerging opportunities and new mechanisms, such as direct and indirect economic incentives for nature conservation and restoration, are taken into account. These, coupled with increased awareness of the stakeholders, better policies and implementation of good governance, and the willingness and know-how to maintain coexistence with wildlife among the local people, are expected to support and accelerate the recovery of tigers and their ecosystem.
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    Ecology of mule deer on the Pinon Canyon Maneuver Site, Colorado
    Gerlach, Thomas P. (Virginia Tech, 1987-01-15)
    Mule deer (Odocoileus hemionus) population dynamics, movements, and habitat use were studied on the Pinon Canyon Maneuver Site in southeastern Colorado during January 1983-December 1984. Thirty-eight adults and 28 fawns were radio collared, and 35 adults were color collared or ear tagged. Population estimates were 365 and 370 deer for 1983 and 1984, respectively. The sex ratio (yearling and adult) was 60 males: 100 females. Adult female pregnancy rate was 95%; the mean litter size for females over 1.5 years was 1.7 fawns. Annual fawn survival was 29% in 1983 and 22% in 1984. Coyote (Canis Iatram) predation was responsible for 76% of fawn mortality. Adult survival was 88% in 1983 and 87% in 1984; coyote predation accounted for 67%, and hunting for 33% of the annual adult mortality. The calculated annual rate of increase (λ) was 1.01, indicating a stable population. Seasonal home range size differed (p < 0.05) between males and females only in the fall. Females preferred pinyon-juniper woodland in all seasons, and shrub grassland in winter, summer and fall; proportional use of woodland/open grassland and shrub/open grassland edge was greater than proportional availability. Males preferred pinyon-juniper woodland and avoided open grassland in all seasons. Fawns preferred shrub grassland and shrub/open grassland edge; they avoided cholla/open grassland edge. Fawns selected bed sites with greater (P < 0.05) concealment cover at all 0.5 m intervals up to 2 m in height, and greater ground cover of trees, shrubs, and grasses (P < 0.01) than random sites.
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    Ecology, Habitat Use, and Conservation of Asiatic Black Bears in the Min Mountains of Sichuan Province, China
    Trent, Jewel Andrew (Virginia Tech, 2010-05-24)
    This project was initiated in an attempt to address the paucity of data on Asiatic black bears (Ursus thibetanus) in Mainland China. Field work was carried out from May 2004 – August 2006 within the Tangjiahe National Nature Reserve in northwestern Sichuan Province, China. Initial methodology relied on trapping and GPS radio-collaring bears, but due to extreme difficulty with capturing a sufficient sample size, I expanded the study to include reproduction, feeding analysis from scats and sign, and occupancy modeling from sign surveys. I documented the home ranges of an adult female (100% MCP = 107.5km2, n=470 locations) and a sub-adult female (100%MCP = 5.9km2, n=36 locations) Asiatic black bear. I also documented two birthing occasions with a total of four male cubs produced and eight bear den sites. I collected feeding data from 131 scat samples and 200 bear sign transects resulting in 50 identified food items consumed by Asiatic black bears. I also employed the program PRESENCE to analyze occupancy data using both a standard grid repeated sampling technique and an innovative technique of aging bear sign along strip transect surveys to represent repeated bear occupancy over time. Conservation protection patrolling and soft mast were shown to be the most important factors determining the occupancy of an area by Asiatic black bears in Tangjiahe Nature Reserve, Sichuan Province, China.
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    The effect of hydroperiod on seed banks in semi-permanent prairie wetlands
    Poiani, Karen A. (Virginia Tech, 1987-06-05)
    In 1985, 24 bottom samples were collected in each of two slightly brackish,semi-permanent prairie wetlands (P1 and P4) with different hydroperiods. The main objective was to determine if hydroperiod affected seed pool characteristics. Additionally, 48 samples were collected in 1986 from wetland P1 to determine if seed bank composition changed annually without a change in mature vegetation. Seed bank composition was determined by placing soil samples in a greenhouse, then counting and identifying emerged seedlings. As a check against the seedling emergence method, seeds were separated and identified microscopically in one-third of the 1985 samples. Results indicated that the emergence method was an accurate technique for assessing seed pool composition. The wetlands did not differ in floristic composition (i.e., presence/absence) but did in species densities. The mean relative density of mudflat annuals in all seed pool samples was significantly greater in wetland P4 (82%) than in P1 (52%). A shorter hydroperiod in this wetland produces more frequent drawdowns and a greater input of mudflat annual seeds. Conversely, seeds of emergent species were more abundant in the seed bank of wetland Pl (48%) compared to P4 (17%). The former wetland has a longer hydroperiod and less frequent drawdowns, and thus, the primary seed input is from emergent plants.
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    Effects of lead ingestion on the immune function in quail
    Grasman, Keith A. (Virginia Tech, 1992-03-05)
    Wild birds can be exposed to high concentrations of lead caused by spent lead shot and industrial waste. The immunosuppressive effects of lead have been demonstrated in laboratory mammals, but little research has been conducted in birds. Because immunocompetence is important for survival, the effects of acute lead exposure on immune function in 2 quail species, Japanese quail (Cotumix coturllix) and northern bobwhites (Colinus virginianus), were investigated. In preliminary experiments, intubation with lead shot produced inconsistent doses within treatment groups. Both quail species exhibited poor antibody responses after immunization with sheep erythrocytes. In a third experiment, male Japanese quail were dosed with 100 or 400 ppm lead in drinking water for 7 days. Quail were fed either laying mash or com. Some quail were treated with corticosterone as a positive control Lead enhanced the loss of body mass caused by com. Lead-induced mortality was observed only in quail fed com. There was marginal evidence that lead increased the heterophil to lymphocyte ratio in quail fed com. Com decreased the cell-mediated immune response to an intradermal injection of phytohemagglutinin. Corticosterone suppressed cell-mediated immunity more than lead. The primary total antibody response to chukar (Alectoris graeca chukar) erythrocytes was significantly suppressed in lead-dosed quail fed com. Lead also reduced the secondary total antibody and IgG responses in the low lead/com group. The most pronounced effect of lead on immune function was a suppression of antibody responses in groups that exhibited clinical lead poisoning.
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    Effects of low level lead and cadmium on reproduction in Peromyscus leucopus and a study of lead concentration in small mammals from old orchards
    Yocum, Susan Marie (Virginia Tech, 1988-03-15)
    Lead and cadmium frequently occur together as contaminants in polluted environments such as roadsides, urban areas, mines, and smelters. Few studies have been conducted to examine the possible interactive effects these metals may have on reproduction of wild species. The first part of this study was directed toward examining the effects of low level dietary lead and cadmium on reproduction in Peromyscus Jeucopus. Two laboratory studies were conducted. Treatment with combinations of 0, 1 or 4 μ/g lead acetate and/or cadmium chloride was found to have no effect on overall health or reproductive ability of white-footed mice. In a second, experiment mice were treated with combinations of 0 or 10 μg/g lead acetate and/or cadmium' chloride. Treatment was continued through three generations of mice (P1, Fl, F2); progeny of the P1 were paired with nonrelatives in the same group and continued on the same treatment. Percentage-of pairs producing at least one litter in a ten week period was reduced in mice receiving both lead and cadmium. By the F2 generation only one of ten pairs produced offspring. There were minor variations in litter parameters (# pups/litter, birth weight, weanling weight and viability of litters) measured for the three generations; however, treatment did not appear to greatly affect litters in those pairs which had litters. In the P1 generation kidney weights were lower in mice receiving lead and epididymidis weight was higher in mice receiving cadmium. Mice (P1) receiving cadmium and lead had lower numbers of spermatozoa/mg testis. Lead content of carcass, liver and kidneys was analyzed in mice from the P1 generation. The second part of this study examined the persistence of lead in the environment after the source of contamination has been removed. Short-tailed shrews (Blarina brevicauda) trapped in an orchard with a history of lead arsenate use had higher whole body lead concentrations than individuals of the same species trapped in a control orchard. Within the treated orchard lead concentrations in pine voles (Microtus pinetorum) were significantly higher than concentrations in short-tailed shrews.
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    Estimating Black Bear Population Size, Growth Rate, and Minimum Viable Population Using Bait Station Surveys and Mark-Recapture Methods
    O'Neill, Deborah M. (Virginia Tech, 2003-09-26)
    We initiated bait station surveys for black bears in southwestern Virginia in 1999. Bait station surveys are intended to be used as an index to follow bear population trend over time. We compared the bait station visitation (black bear visitation) to black bear harvest and mast surveys 1999 = 2002. The mean bait station visitation rate during 1999 - 2002 was 15.3% (SE = 2.89, n = 4). The number of bears harvested in the 3 counties that also had bait station surveys was 48 (31 males, 17 females), 59 (44 males, 15 females), 45 (32 males, 13 females), and 43 (26 males, 17 females) in 1999, 2000, 2001, and 2002, respectively. Harvest of males and females differed (n = 2, F = 19.44, df = 1, P = 0.0045). Bait station visitation and female harvest had a strong functional relationship with a negative slope (n = 4, r = -0.78, P = 0.22). The strongest relationship was between male harvest and total harvest (n = 4, r = 0.97, P = 0.03). Mean index to mast production for 1999 - 2002 was 2.3 (range 1.5 - 3.1), 2.7 (range 1.8 - 3.4), 2.3 (range 1.6 - 3.6), and 1.6 (range 1.2 - 2.4), respectively. The overall summary for mast production for the same years was described as fair, good, fair, and poor to fair. Mast production was significantly different between years (n = 4, F = 3.44, df = 3, P = 0.0326), and soft and hard mast production appeared to be above average in 2000. This corresponded with the lowest visitation (10.2%) of the 4 years. There was no correlation between bait station visitation and mast production (n = 4, r = 0.11, P = 0.87). Since 1998, the annual bear harvest in Virginia has exceeded 900 individuals (with the exception of 824 in 2001), and peaked in 2000 when 1,000 bears were harvested. Though harvest rates were high, a reliable population estimate did not exist for black bears in Virginia. We estimated population size, growth rate, and minimum viable population size using data collected between 1995-2000. We used Jolly-Seber, direct recovery, and minimum population size methods to estimate population size. The Jolly-Seber method estimate of adult female density was 0.23-0.64 bears/km2, and 0.01 bears/km2 for adult males. We estimated a density of 0.09-0.23 bears/km2 for all sex and age classes using direct recovery data. Using minimum population size, we found adult female density was higher than any other sex or age class (n = 6, t = 2.02, df = 40, P < 0.0001) with an average density of 0.055 adult females/km2. We used mark-recapture data collected from 148 individual bears (96 males:52 females) captured 270 times in program MARK to estimate survival using recapture, dead recovery, and Burnham's combined models. Adult females had the highest survival rate of 0.84-0.86, while yearling males had the lowest with 0.35. Using direct recovery data, adult females again had the highest survival rate with 0.93 (0.83-1.0) and 3-year old males had the lowest with 0.59 (0.35-0.83). We estimated growth rate using population estimates from Jolly-Seber, direct recoveries, and minimum population size methods. The lowest growth rate estimated was for all females (ages lumped) using minimum population size data (λ=0.82). Direct recovery data for all bears (sex and age lumped) during 1995 - 2000 showed the highest positive annual growth rate (λ = 1.24). We developed a population model using Mathcad 8 Professional to determine population growth rate, MVP, and harvest effects for an exploited black bear population in southwestern Virginia. We used data collected during the CABS study (1995 - 2000) in the model including population estimates derived from direct recovery data, age and sex specific survival rates, and cub sex ratios. When we used actual population values in the model, the bear population in southwestern Virginia did not go extinct in 100 years (l = 1.03, r = 0.03). When we reduced adult female survival from 0.94 to 0.89, the probability of extinction in 100 years was 3.0% and l = 0.99 (r = -0.01; Table 3.2). When the survival was reduced by an additional 0.01 to 0.88, the probability of extinction increased to 13.0% (l = 0.99, r = -0.01). Growth rate and extinction probabilities were very sensitive to adult female survival rates. Two-year old and 3-year old females did not impact extinction probabilities and growth rates as much as adult females. Their survival could be decreased by 44.0%, and still be less than the 5.0% extinction probability.
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    Evaluating the Effectiveness of Population Reconstruction for Black Bear (Ursus americanus) and White-Tailed Deer (Odocoileus virginianus) Population Management
    Tilton, Mary Kathryn (Virginia Tech, 2005-09-14)
    This study was a comprehensive evaluation of population reconstruction techniques. Population reconstruction techniques are population estimation methods that calculate a minimum population size based on age-specific harvest data (Downing 1980, Roseberry and Woolf 1991). Population reconstruction techniques share the following characteristics: 1) utilization of catch-at-age data and 2) backward addition of cohorts to estimate a minimum population size. I developed a questionnaire to survey the biologists participating in this survey to determine the most common reconstruction technique used to estimate population sizes of exploited white-tailed deer (Odocoileus virginianus) and black bear (Ursus americanus). Downing reconstruction (Downing 1980) was the most commonly used reconstruction technique among biologists participating in this study. Based on a comprehensive literature review and discussions with state biologists, I decided to evaluate virtual reconstruction (Roseberry and Woolf 1991) and develop a new reconstruction technique: Reverse Order reconstruction. I developed a quantitative population model in Microsoft Visual Basic 6.0 to evaluate the ability of the 3 reconstruction techniques to estimate population sizes given a variety of conditions. I evaluated the effects of stochasticity on reconstruction population estimates by incorporating different levels of environmental stochasticity (i.e. process error) and measurement error in the generated or "known" population. I also evaluated the effects of collapsing age classes and aging biases on population estimates. In all conditions, Downing and virtual reconstruction were underestimates of the actual population size. Reverse Order reconstruction more closely estimated the actual population size, but is also more data-intensive than the other 2 methods. Measurement error introduces more uncertainty in the reconstructed population estimates than does process error. The population simulation model proved that Downing and virtual reconstruction are consistently underestimates and the percent underestimation is due to lack of inclusion of a natural mortality rates in population estimation. I used the results of the questionnaire to characterize the harvest datasets of the states participating in this study. From these results, I chose two harvest datasets to further analyze: a white-tailed deer harvest dataset from North Carolina and a black bear harvest dataset from Pennsylvania. I analyzed these datasets with Downing and virtual reconstruction. I also applied the quantitative population model to these datasets to evaluate the effect of increasing levels of measurement error on the variance of the population estimates. I found that Downing and virtual reconstruction estimated the population sizes very closely to one another, within 5%, for both datasets, and the reconstructed estimates closely tracked the actual harvest numbers. I also found that increasing levels of measurement error increased the variance associated with reconstructed population estimates and may decrease the ability of these techniques to accurately capture population trends.
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    Evolution of stakeholder knowledge, attitudes, and opinions throughout a participative process to develop a management plan for black bears in Virginia
    Lafon, Nelson Wade (Virginia Tech, 2002-02-01)
    Wildlife management requires understanding resources and involving stakeholders. The process Virginia Department of Game and Inland Fisheries (VDGIF) and Virginia Tech used to develop a black bear (Ursus americanus) management plan during 1999-2001 provided me with opportunities to understand diverse stakeholders and examine influences of participation on stakeholder knowledge, attitudes, and opinions concerning bear management. I used focus groups (with 5 key stakeholder groups), pre- and post-planning surveys, and interviews. I surveyed VDGIF biologists and managers (N = 21), members of a stakeholders advisory committee (N = 15), and members of 3 constituent groups representing bear hunters (N = 459), beekeepers (N = 442), and environmental interests (N = 500). I interviewed advisory committee members to validate survey results and evaluate the bear management plan. Participation apparently improved constituents' knowledge about bear management and their image of VDGIF bear management, and increased their support for controversial management options (e.g., lethal methods), but did not affect their opinions about bear hunting. Participation apparently had little effect on VDGIF staff opinions about bear management. Knowledge varied widely among constituent organizations. Bear hunting continues to be one of the most central, yet divisive, bear management issues in Virginia. Advisory committee members and VDGIF staff expressed satisfaction with the bear management plan and the planning process. Recommendations for wildlife decision-making processes include: balance science and public values in management, use multiple public involvement techniques, establish collaborative forums among stakeholders, reach out to all stakeholders, and nurture relationships with constituents during implementation.
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    Factors Influencing Predation on Ruffed Grouse in the Appalachians
    Bumann, George Bruce (Virginia Tech, 2002-04-12)
    Predation accounts for the majority of ruffed grouse mortality beyond the brooding period throughout grouse range. Most studies of ruffed grouse ecology have taken place in the central portion of the species range including the Great Lakes states and southern Canada where aspen (Populus tremuloides) is dominant. Populations in the central range of grouse distribution exhibit 10-year cyclic patterns of decline that have been associated with the invasions of boreal predators such as great horned owls and northern goshawks during crashes in small mammal populations. I completed an accuracy assessment of field sign to determine the role of scavenging as it related to diagnosing causes of proximate mortality. Potentially large numbers of grouse are scavenged after death, which can alter or completely falsify the reported cause of death.. My results indicated that stand-level and micro-site habitat variables did not influence scavenger detection of ruffed grouse carcasses (P > 0.05). Scavenging was limited to 24 of 64 carcasses and was entirely attributed to mammalian species. Scavenging behavior was related to the condition of the carcass following death (P = 0.003) and the ambient temperature (P = 0.01). As the temperature increased and as a carcasses entrails and muscle became more exposed, the probability of being scavenged increased. Nineteen percent of whole carcasses placed in the field were scavenged and would have been attributed to mammal predation based on the field sign; 56% of 32 mock avian kills were scavenged would have been attributed to mammal predation. I related indices of predators and weather patterns to trends in ruffed grouse predation. Using data collected at 10 study sites between February, 1997 and December, 2000, I compared predation rates, and animal and weather indices to predation rates, across sites, years, regions, seasons and month combinations pooled across years. Avian predators were the primary predators of ruffed grouse in the Appalachian region (50% of all predation). Predation rates on ruffed grouse were highest in fall (8.3%), due to high predation on juveniles, and spring (7.4%) in association with raptor migration and pre-breeding activity of grouse. Predation patterns and predator abundance did not indicate the occurrence of predator invasions during the years of this study. Predation rates on ruffed grouse were positively related to the presence of rain and negatively related to the average low temperature and number of rabbits and squirrels observed per hour. Observations of owls and Cooper's hawks per hour were correlated with predation rates on grouse while those of red-tailed, red-shouldered and broad-winged hawks were not.