Joseph D. Clark

A Multivariate Model of Female Black Bear Habitat Use for a Geographic Information System

Simple univariate statistical techniques may not adequately assess the multidimensional nature of habitats used by wildlife. Thus, we developed a multivariate method to model habitat-use potential using a set of female black bear (Ursus americanus) radio locations and habitat data consisting of forest cover type, elevation, slope, aspect, distance to roads, distance to streams, and forest cover type diversity score in the Ozark Mountains of Arkansas. The model is based on the Mahalanobis distance statistic coupled with Geographic Information System (GIS) technology. That statistic is a measure of dissimilarity and represents a standardized squared distance between a set of sample variates and an ideal based on the mean of variates associated with animal observations

Spatially explicit integrated population models

Summary Studies of demographic processes are typically restricted to small geographic areas and short time periods due to the costs of marking and monitoring individuals. However, environmental changes are occurring at much broader spatial and temporal scales, and thus, inferences about the mechanisms governing population dynamics need to be scaled accordingly. Recently developed integrated population models (IPMs) represent an approach for doing so, by jointly analysing survey data and capture–recapture data. Although promising, several shortcomings of conventional IPMs exist, including difficulties accounting for spatial variation in demographic, movement and detection parameters; limited ability to make spatially explicit predictions of abundance or vital rates; and a requirement that the survey data and the capture–recapture data are independent. We demonstrate how each of these limitations can be resolved by adopting a spatial population dynamics model upon which both the survey data and the capture–recapture data are conditioned. We applied the model to 6 years of hair data collected on the threatened Louisiana black bear Ursus americanus luteolus. For years in which the hair samples were genotyped, the resulting data are information-rich (but expensive) spatial capture–recapture (SCR) data. For the remaining years, the data are binary detection data, of the type often analysed using occupancy models. We compared estimates of demographic parameters and annual abundance using various combinations of the SCR and detection data, and found that combining the SCR data and the detection data resulted in more precise estimates of abundance relative to estimates that did not use the detection data. A simulation study provided additional evidence of increased precision, as well as evidence that the estimators of annual abundance are approximately unbiased. The ability to combine survey data and capture–recapture data using a spatially explicit model opens many possibilities for designing cost effective studies and scaling up inferences about the demographic processes influencing spatial and temporal population dynamics.

Ecology of Florida Black Bears in the Okefenokee‐Osceola Ecosystem

Abstract: The population status of the Florida black bear (Ursus americanus floridanus) is problematic within many portions of its range and its potential listing as a federally threatened species has been the subject of legal debate. We studied Florida black bears in 2 areas in the Okefenokee-Osceola ecosystem in southeast Georgia (i.e., Okefenokee) and north Florida (i.e., Osceola) from 1995 to 1999 to evaluate relationships between population characteristics, habitat conditions, and human activities. Bears in Okefenokee were hunted and those in Osceola were not. We captured 205 different black bears (124M:81F) 345 times from June 1995 to September 1998. We obtained 13,573 radiolocations from 87 (16M:71F) individual bears during the study. In Okefenokee, black gum (Nyssa sylvatica) and saw palmetto (Serenoa repens) fruits were the most important foods for bears based on scat analysis. In Osceola, corn from white-tailed deer (Odocoileus virginianus) feeders was the most stable food source but saw palmetto was heavily used when available. Corn from deer feeders was not available in Okefenokee. Adult bears in Osceola were 29% heavier than those in Okefenokee (t82 = 3.55, P < 0.001). The mean annual home-range size for Osceola females (x = 30.3 km2 ± 4.0 [SE], n = 53) varied little seasonally or annually and was almost half that of Okefenokee females (55.9 km2 ± 6.9, n = 69; Z = −2.47, P = 0.014). In contrast, radiocollared females in Okefenokee expanded their home ranges during years of poor black gum production. That expansion was most apparent between autumn 1998 and 1999, when mean home-range size for Okefenokee females increased from 14.5 km2 to 78.4 km2, respectively, and included a larger proportion of upland areas open to sport hunting. As a result, 5 females were harvested in the Okefenokee study area during the 1999 bear hunting season compared with only 7 harvested from 1996 to 1998. Home ranges of adult female bears were located in areas with disproportionately high loblolly bay (Gordonia lasianthus) and gum-bay-cypress (Taxodium spp.) vegetation associations in Okefenokee and gum-bay-cypress associations in Osceola. The pine vegetation association ranked lower than most other associations within the home ranges of bears in both study areas even though much of the summer and autumn diets of bears included food items found almost exclusively in pine. Sixteen mortalities of radiocollared bears were documented in Okefenokee; hunting accounted for 11 (68.8%) of these deaths. The annual survival rate of radiocollared males in Okefenokee was 0.71 (95% CI = 0.53-0.88) whereas survival of females in Okefenokee was higher (Z = 18.87, P < 0.001) at 0.89 (95% CI = 0.83-0.95). The survival rate for females in Osceola was 0.97 (95% CI = 0.92-1.00). Overall, 67 bears (51M:16F) were killed by hunters in the Okefenokee study area from 1995 to 1999. Based on all radiocollared bears in Okefenokee, many of which resided within areas closed to hunting, we estimated an annual harvest rate of 0.22 (95% CI = 0.03-0.37) for males and 0.07 (95% CI = 0.01-0.12) for females. When we excluded those bears that were not in areas open to hunting, however, the annual harvest rate rose to 0.37 (95% CI = 0.07-0.58) for males and 0.39 (95% CI = 0.09-0.58) for females. Following a black gum shortage in autumn 1995, only 1 of 15 radiocollared females in Okefenokee produced cubs in winter 1996. That low reproductive rate was in contrast to winter 1997, which followed heavy black gum production, when 21 of 22 radiocollared females produced cubs. Reproductive output was more consistent in the Osceola study area, with 46 cubs being produced from 8, 5, and 9 litters in 1997, 1998, and 1999, respectively. To estimate population size, we maintained 88 and 94 barbed-wire hair traps during 1999 in the Okefenokee and Osceola study areas, respectively. Using DNA collected at the hair traps, mark-recapture models produced estimates of 71 bears (95% CI = 59–91) in Okefenokee and 44 bears (95% CI = 40–57) in the Osceola study area during 1999. The estimated densities in the Okefenokee and Osceola study areas were 0.12 and 0.14 bears/km2, respectively. Alternative density estimates based on the amount of time bears spent within study area boundaries were similar (0.11 and 0.14 bears/km2 on Okefenokee and Osceola, respectively). We used a population model to estimate the effect of harvest in the Okefenokee bear population. Excluding harvest, bears at Osceola experienced higher average annual population growth (Λ = 1.184 ± 0.002) than those at Okefenokee (1.064 ± 0.002; t18 = 3.93, P = 0.001), most likely due to protection from hunting and higher recruitment. Including the effects of emigration and immigration, we estimated an average annual sustainable harvest at Okefenokee of approximately 9 bears (12.6%), which was slightly less than the average 1995–1999 annual harvest of 9.4. That level of hunting in Okefenokee is sustainable but likely represents the highest exploitation rate in the region. Our mark-recapture data from Osceola suggest a high dispersal rate by subadult bears, and our population modeling data support this hypothesis; we documented bears in Okefenokee that originated from Osceola but not the reverse. We speculate that bears from the interior of the Okefenokee National Wildlife Refuge (ONWR), and to some extent northern Florida, served as a source to the population sink caused by hunting mortality in Okefenokee and in the surrounding Georgia counties. Corn from deer feeders was the most probable reason for smaller home-range sizes and greater body masses and reproductive output at Osceola. Changes in management to eliminate or reduce baiting for deer with corn would negatively affect the Osceola bear population. On Okefenokee, sporadic black gum and palmetto production influenced harvest rates and cub production and, thus, governed bear population dynamics. The U.S. Fish and Wildlife Service (USFWS) concluded in 1998 that listing the Florida black bear as threatened under the Endangered Species Act of 1973 was not warranted. That decision was largely based on the stability and protection afforded to a few subpopulations within the range of the subspecies, which includes the Okefenokee-Osceola subpopulation; our results support that conclusion. However, we suggest that metapopulation processes among the various subpopulations be given greater consideration, with the ultimate goal of managing the subspecies as a unit rather than as an assemblage of independent components. Our study illustrates the importance of travel corridors for maintaining metapopulation processes.

Identifying Suitable Sites for Florida Panther Reintroduction

Abstract A major objective of the 1995 Florida Panther (Puma concolor coryi) Recovery Plan is the establishment of 2 additional panther populations within the historic range. Our goal was to identify prospective sites for Florida panther reintroduction within the historic range based on quantitative landscape assessments. First, we delineated 86 panther home ranges using telemetry data collected from 1981 to 2001 in south Florida to develop a Mahalanobis distance (D2) habitat model, using 4 anthropogenic variables and 3 landscape variables mapped at a 500-m resolution. From that analysis, we identified 9 potential reintroduction sites of sufficient size to support a panther population. We then developed a similar D2 model at a higher spatial resolution to quantify the area of favorable panther habitat at each site. To address potential for the population to expand, we calculated the amount of favorable habitat adjacent to each prospective reintroduction site within a range of dispersal distances of female panthers. We then added those totals to the contiguous patches to estimate the total amount of effective panther habitat at each site. Finally, we developed an expert-assisted model to rank and incorporate potentially important habitat variables that were not appropriate for our empirical analysis (e.g., area of public lands, livestock density). Anthropogenic factors heavily influenced both the landscape and the expert-assisted models. Of the 9 areas we identified, the Okefenokee National Wildlife Refuge, Ozark National Forest, and Felsenthal National Wildlife Refuge regions had the highest combination of effective habitat area and expert opinion scores. Sensitivity analyses indicated that variability among key model parameters did not affect the high ranking of those sites. Those sites should be considered as starting points for the field evaluation of potential reintroduction sites.

Florida panther habitat use in response to prescribed fire

The Florida panther (Puma concolor coryi) is one of the most endangered mammals in the world, with only 30-50 adults surviving in and around Florida Panther National Wildlife Refuge and the adjacent Big Cypress National Preserve. Managers at these areas conduct annual prescribed burns in pine (Pinus sp.) as a cost-effective method of managing wildlife habitat. Our objectives were to determine if temporal and spatial relationships existed between prescribed fire and panther use of pine. To accomplish this, we paired fire-event data from the Refuge and the Preserve with panther radiolocations collected between 1989 and 1998, determined the time that had elapsed since burning had occurred in management units associated with the radiolocations, and generated a frequency distribution based on those times. We then generated an expected frequency distribution, based on random use relative to time since burning. This analysis revealed that panther use of burned pine habitats was greatest during the first year after a management unit was burned. Also, compositional analysis indicated that panthers were more likely to position their home ranges in areas that contained pine. We conclude that prescribed burning is important to panther ecology. We suggest that panthers were attracted to <1-year-old burns because of white-tailed deer (Odocoileus virginianus) and other prey responses to vegetation and structural changes caused by the prescribed fires. The strong selection for stands burned within 1 year is a persuasive indication that it is the burning in pine, rather than the pine per se, that most influenced habitat use. Before burning rotation lengths are reduced, however, we suggest managers determine effects of shorter burning intervals on vegetation composition and evaluate the landscape-scale changes that would result.

Responses of Florida panthers to recreational deer and hog hunting

Big Cypress National Preserve constitutes approximately one-third of the range of the endangered Florida panther (Puma concolor coryi). Because recreational hunting is allowed in Big Cypress National Preserve, we examined 8 response variables (activity rates, movement rates, predation success, home-range size, home-range shifts, proximity to off-road vehicle trails, use of areas with concentrated human activity, and habitat selection) to evaluate how Florida panthers respond to human activity associated with deer and hog hunting. Data consisted of panther radiolocations collected since 1981 by the Florida Fish and Wildlife Conservation Commission and the National Park Service, which we augmented with radiolocations and activity monitoring from 1994 to 1998. A split-plot (treatment and control) study design with repeated measures of the variables for each panther taken before, during, and after the hunting season was used. We did not detect responses to hunting for variables most directly related to panther energy intake or expenditure (i.e., activity rates, movement rates, predation success of females; P>0.10). However, panthers reduced their use of Bear Island (P = 0.021), an area of concentrated human activity, and were found farther from off-road vehicle trails (P ≤ 0.001) during the hunting season, which was indicative of a reaction to human disturbance. Whereas the reaction to human activity on off-road vehicle trails probably has minor biological implications and may be linked to prey behavior, the decreased use of Bear Island is most likely a direct reaction to human activity and resulted in increased use of adjacent private lands. Future habitat loss on those private lands could exacerbate the negative consequences of this response by panthers.

GENETIC VARIATION IN BLACK BEARS IN ARKANSAS AND LOUISIANA USING MICROSATELLITE DNA MARKERS

Abstract In the 1950s and 1960s, translocation projects reintroduced black bears (Ursus americanus) from Minnesota and Manitoba to Arkansas and Louisiana. Today, several geographically disconnected populations exist in Arkansas and Louisiana, but their origins are unclear. Some populations may represent a separate subspecies, U. a. luteolus, which is federally protected. We characterized 5 microsatellite loci in 5 isolated populations in Arkansas and Louisiana and compared them with genotypes from Minnesota. Our data indicate that bears of the Ozark and Ouachita mountains of Arkansas, an inland area of Louisiana, and those of Minnesota are similar in overall genetic diversity and allele frequencies, consistent with these populations being wholly or mostly descended from bears from the reintroduction programs. In contrast, bears from southeastern Arkansas and the coastal region of Louisiana genetically are more restricted and homogeneous. Because they exhibit a limited set of genotypes found in the other black bear populations, they represent isolated fragments of a single North American black bear population. Furthermore, genetic distance estimates indicate that the bears in southeastern Arkansas are more genetically distinct from bears in Louisiana, which are currently federally protected.

Challenges of DNA-based mark-recapture studies of American black bears

Abstract We explored whether genetic sampling would be feasible to provide a region-wide population estimate for American black bears (Ursus americanus) in the southern Appalachians, USA. Specifically, we determined whether adequate capture probabilities (p > 0.20) and population estimates with a low coefficient of variation (CV < 20%) could be achieved given typical agency budget and personnel constraints. We extracted DNA from hair collected from baited barbed-wire enclosures sampled over a 10-week period on 2 study areas: a high-density black bear population in a portion of Great Smoky Mountains National Park and a lower density population on National Forest lands in North Carolina, South Carolina, and Georgia. We identified individual bears by their unique genotypes obtained from 9 microsatellite loci. We sampled 129 and 60 different bears in the National Park and National Forest study areas, respectively, and applied closed mark–recapture models to estimate population abundance. Capture probabilities and precision of the population estimates were acceptable only for sampling scenarios for which we pooled weekly sampling periods. We detected capture heterogeneity biases, probably because of inadequate spatial coverage by the hair-trapping grid. The logistical challenges of establishing and checking a sufficiently high density of hair traps make DNA-based estimates of black bears impractical for the southern Appalachian region. Alternatives are to estimate population size for smaller areas, estimate population growth rates or survival using mark–recapture methods, or use independent marking and recapturing techniques to reduce capture heterogeneity.

Growth and Sustainability of Black Bears at White River National Wildlife Refuge, Arkansas

Abstract The black bear (Ursus americanus) population at White River National Wildlife Refuge is isolated and genetically distinct, but hunting occurs adjacent to refuge boundaries and females with cubs are removed annually for a reintroduction project. We trapped and radiotracked bears to determine level of exploitation and compare methods for estimating population growth and sustainability. We captured 260 bears (113 M:147 F), 414 times, from 1998 through 2003. Survival estimates based on radiotracking and mark–recapture indicated that hunting and translocations were significant sources of loss. Based on mark–recapture data (Pradel estimator), the annual population growth rate (λ) averaged 1.066 (SE = 0.077) when translocation removals occurred and averaged 0.961 (SE = 0.155) when both harvest and translocations occurred. Estimates of λ based on a population simulation model (program RISKMAN) averaged 1.061 (SD = 0.104) and 1.100 (SD = 0.111) when no removals occurred, 1.003 (SD = 0.097) and 1.046 (SD = 0.102) when translocations occurred, and 0.973 (SD = 0.096) and 1.006 (SD = 0.099) when both harvest and translocations occurred, depending on the survival rate estimates we used. The probability of population decline by >25% over a 10-year period ranged from 13.8 to 68.8%, given our estimated removal rates. We conclude that hunting and translocation losses are at or exceed the maximum the population is capable of sustaining. Although extinction risks of this important bear population are low over the near term, it should continue to be closely monitored by state and federal agencies. The mark–recapture method we used to estimate λ proved to be a reliable alternative to more costly population modeling methods.

Factors affecting settling, survival, and viability of black bears reintroduced to Felsenthal National Wildlife Refuge, Arkansas

Abstract We used radiotelemetry and population modeling techniques to examine factors related to population establishment of black bears (Ursus americanus) reintroduced to Felsenthal National Wildlife Refuge (NWR), Arkansas. Our objectives were to determine whether settling (i.e., establishment of a home range at or near the release site), survival, recruitment, and population viability were related to age class of reintroduced bears, presence of cubs, time since release, or number of translocated animals. We removed 23 adult female black bears with 56 cubs from their winter dens at White River NWR and transported them 160 km to man-made den structures at Felsenthal NWR during spring 2000–2002. Total movement and average circuity of adult females decreased from 1 month, 6 months, and 1 year post-emergence (F2, 14 =19.7, P <0.001 and F2, 14 =5.76, P =0.015, respectively). Mean first-year post-release survival of adult female bears was 0.624 (SE=0.110, SEinterannual= 0.144), and the survival rate of their cubs was 0.750 (SE=0.088, SEinterannual=0.109). The homing rate (i.e., the proportion of bears that returned to White River NWR) was 13%. Annual survival for female bears that remained at the release site and survived >1-year post-release increased to 0.909 (SE=0.097, SEinterannual=0.067; Z=3.5, P<0.001). Based on stochastic population growth simulations, the average annual growth rate (λ) was 1.093 (SD=0.053) and the probability of extinction with no additional stockings ranged from 0.56–1.30%. The bear population at Felsenthal NWR is at or above the number after which extinction risk declines dramatically, although additional releases of bears could significantly decrease time to population reestablishment. Poaching accounted for at least 3 of the 8 adult mortalities that we documented; illegal kills could be a significant impediment to population re-establishment at Felsenthal NWR should poaching rates escalate.