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37 $QWKURSRJHQLF,QÀXHQFHRQ$PHULFDQ%ODFN%HDU 'LHWLQWKH:HVWHUQ2]DUN0RXQWDLQVLQ(DVWHUQ 2NODKRPD Joseph P. Connor School of Natural Sciences, University of Central Missouri, Warrensburg, MO 64093 9LFWRULD/-DFNVRQ Department of Biology, University of Central Oklahoma, Edmond, OK 73034 -HQQLIHU50LWWHOKDXVHU School of Natural Sciences, University of Central Missouri, Warrensburg, MO 64093 :6XH)DLUEDQNV Department of Natural Resource Ecology and Management, Oklahoma State University, Stillwater, OK 74078

$EVWUDFW American black ( americanus) are returning to eastern Oklahoma from Arkansas, where they were re-introduced in the 1950s. This movement back into -occupied DUHDVFDQOHDGWRFRQÀLFW%ODFNEHDUVDUHNQRZQWRXVHDQWKURSRJHQLFIRRGVRXUFHV7RGHWHUPLQH WKHH[WHQWRIDQWKURSRJHQLFLQÀXHQFHZHDQDO\]HGVFDWIRUDQWKURSRJHQLFIRRGLWHPV:HFROOHFWHG  VFDW VDPSOHV IURP  0D\  WKURXJK  2FWREHU  LQ WKH 2]DUN 3ODWHDX RI $GDLU Cherokee, and Sequoyah counties. Once collected, scat was dried for preservation. The samples were UHK\GUDWHGDQG¿OWHUHGIRUXQGLJHVWHGLWHPVZKLFKZHUHFDWHJRUL]HGLQWRJURXSV:HPHDVXUHG percent volume and percent frequency of occurrence of each item that was > 1% of the volume of the HQWLUHVDPSOH7KHYROXPHVRIGLHWJURXSVZHUHVLJQL¿FDQWO\GL൵HUHQW .UXVNDO:DOOLVS   and were separated into 3 statistically distinct groups: Anthropogenic food > Reproductive plant SDUWV!$QLPDOPDWWHU+HUEDJH'HEULVDQG8QLGHQWL¿HG 7XNH\¶V+RQHVW6LJQL¿FDQW'L൵HUHQFH WHVWDOSKD  7KHUHZDVQRVLJQL¿FDQWGL൵HUHQFHEHWZHHQWKHYROXPHVRIDQWKURSRJHQLFYHUVXV natural food (Mann Whitney U, p = 0.368). Though black bears will eat anthropogenic food sources LIWKH\HQFRXQWHUWKHPDQWKURSRJHQLFIRRGVRXUFHVGRQRWPDNHXSDVLJQL¿FDQWSRUWLRQRIWKHLUGLHW when compared against all natural forage.

preference (Breck et al. 2009).

,QWURGXFWLRQ )RRG DEXQGDQFH DQG GLVWULEXWLRQ FDQ D൵HFW movements and behavior of (Isbell +XPDQDFWLYLW\FDQD൵HFWZLOGOLIHSRSXODWLRQV et al. 1998), and often more so in carnivores in negative ways such as transportation of (Jepsen et al. 2002). For some species, such as invasive species, over-harvest of species, habitat the (Procyon lotor PDQ\RIWKHH൵HFWV loss, and habitat fragmentation (Forester and RIDQWKURSRJHQLFLQÀXHQFHDUHSRVLWLYHVXFKDV 0DFKOLV  :RRGUR൵H  0F.LQQH\ increased food supply through anthropogenic    )XUWKHUPRUH KXPDQ LQÀXHQFH LV PHDQVDQGDGHFUHDVHLQQDWXUDOSUHGDWRUV 3UDQJH FKDQJLQJVSHFL¿FDQLPDOEHKDYLRUVVXFKDVGLHW et al. 2003). For other species, such as American Proc. Okla. Acad. Sci. 98: pp 37 - 45 (2018) 38 J.P. Connor, V.L. Jackson, J.R. Mittelhauser, and W.S. Fairbanks black bears (Ursus americanus), anthropogenic  &RQÀLFWVPLJKWDOVRLQFOXGHSUHGDWLRQRQ LQÀXHQFHLVDOPRVWDOZD\VQHJDWLYHPDLQO\GXH livestock, a problem that is on the rise in Colorado to the fragmentation of habitat and human- (Baruch-Mordo et al. 2008). Based on reports FRQÀLFW 0DWWVRQ  . &RQÀLFW RFFXUV ZKHQ of depredation to the Colorado Department EHDUVWU\WRXWLOL]HDQWKURSRJHQLFIRRGVRXUFHV of Wildlife from 1979 to 2003, complaints of such as trash containers, gardens, orchards, depredation by bears mostly involved sheep, but apiaries, and corn (Merkle et al. 2013). This also included goats, chickens, pigs, and cows problem is exacerbated when purposely (Baruch-Mordo at al. 2008). Bears are more provide anthropogenic food sources to wildlife, likely to seek out anthropogenic food sources such as deer feeders. when there is a shortage of natural forage (Clark et al. 2005). In West Virginia, from 1980-2004, Understanding how wildlife populations black bear mortality increased in years when H[SDQGWRUHFRORQL]HIRUPHUKDELWDWVLVLPSRUWDQW there was mast failure. These deaths were mainly for management and conservation (Swenson et due to road kills and landowners destroying al. 1998). American black bears can be found in bears that damaged property (Ryan et al. 2007). relative abundance within much of the northern U.S. but are rare in the southern states. However, American black bears require a variety of EHDUV DUH EHJLQQLQJ WR UHFRORQL]H VRXWKHUQ habitats that produce seasonal foods, as well areas in the U.S., including Oklahoma. Black as extensive and secluded areas for denning bears were extirpated from Oklahoma by 1915 (Landers et al. 1979). Habitat selection by bears but they were reintroduced in Arkansas in the varies seasonally and is governed by presence 1950s and 1960s (Smith and Clark 1994). The of food (Clark et al. 1994; Fuller and Keith population has expanded back into Oklahoma 1980). Though they are part of Order , and southern Missouri (Bales et al. 2005). The black bears are omnivorous. These southern population in Oklahoma, found in the prefer heavily wooded areas with mast species Ouachita Mountains, is well established and like oak (Quercus spp.), hickory (Carya spp.), JURZLQJ 3IDQGHU ,QWKH2NODKRPD and various species of berries (Benson and Department of Wildlife Conservation opened a Chamberlain 2006). Black bears rarely hunt black bear hunting season for the counties of but have been known to take neonate ungulates /H)ORUH0F&XUWDLQ3XVKPDWDKDDQG/DWLPHU (Schlegel 1976). When available, they will also The east-central population in Adair, Cherokee, eat carrion (Arner 1948). and Sequoyah counties is not yet self-sustaining and is not hunted (Lyda et al. 2016). In the fall, black bears prefer a greater proportion of natural foods due to the The east-central black bear population is availability of hard mast such as acorns, hickory UHFRORQL]LQJ LQ DQ DUHD WKDW DOUHDG\ KDV VRPH nuts, walnuts (Juglans spp.) and beechnuts human habitation, with 22,098 people in Adair (Fagus spp.) (Sara Lyda, [Oklahoma State County (14.8 people/km2), 48,700 people in University, Stillwater, Oklahoma], personal Cherokee County (24.2 people/km2), and 41,294 communication, [September 2013]). The hard people in Sequoyah County (22.3 people/km2) mast species are still important in the spring (U.S. Census Bureau 2016). As these animals when females emerge from their dens with cubs. move in, they may begin to use anthropogenic Quantity and quality of acorns determines the food sources such as apiaries, deer feeders, quality and quantity of milk produced by sows orchards, and trash bins (Merkle et al. 2013). (McDonald and Fuller 2005). Milk is high in fat In Oklahoma, it is legal to bait deer and other (220 g/kg) and low in water (670 g/kg), helping wildlife on private land. Availability of this bait, altricial cubs to gain weight quickly during XVXDOO\FRUQPD\KDYHDPDMRULQÀXHQFHRQWKH nursing (Oftedal et al. 1993). overall diet of the bears and other wildlife in the area. Bears have been known to break into Our research addressed the proportion of cars if they think food is inside (Breck et al. bear diet comprised of anthropogenic resources Proc. Okla. Acad. Sci. 98: pp 37 - 45 (2018) Black Bear Diet in Oklahoma 39

LQWKH2]DUN3ODWHDXRIHDVWHUQ2NODKRPD:H VLWHV $OO VDPSOHV ZHUH SODFHG LQWR ]LSORFN also determined the general composition of bags for storage. We recorded information the diet based on scat analysis. Because of the about the scat such as: date collected, collector availability of deer feeders, we predicted that name, sample ID number, whether the sample anthropogenic foods would account for the was whole or partial, estimate of sample age, greatest volume compared to natural foods. sample color, distance to anthropogenic food source, visible solid matter, UTM or latitude and 0HWKRGV longitude, county of collection, canopy cover, and a description of the collection site. Samples As part of a larger project addressing were considered partial if only a small amount population demography, both live capture and of scat was collected or if the sample collected non-invasive genetic methods were used to consisted of 2 or more scats and could not be identify individual bears. This project followed separated. The partial samples were not included the American Society of Mammalogists (ASM) in the volume and frequency analysis because guidelines for the humane use of mammals they do not represent a single whole sample. 6LNHVHWDO DQGZDVDSSURYHGE\268¶V Samples were removed from bags later that day Institutional Care and Use Committee and dried by heat lamp to preserve for future ,$&8& XQGHU3URWRFRO$* analysis. In October 2014, we began rehydrating the samples. Samples were placed into a rubber 7KHDUHDRIVWXG\ZDVWKH2]DUN3ODWHDXLQ tub with enough water and Dawn™ dish soap Adair, Cherokee, and Sequoyah counties in to submerge the samples and then were left to Oklahoma. From 1 May 2014 to 1 September VRDN IRU DSSUR[LPDWHO\  KRXU *UDEHU  2014, average temperature for Cookson, OK, *UHHQOHDIHWDO 7KH'DZQŒGLVKVRDS ZDVÛ& Û) ZLWKDUDQJHRI served as a surfactant to lower the surface Û&DQGWKHDYHUDJHSUHFLSLWDWLRQZDVFP tension of the water. This was important to not per day (Oklahoma Climatological Survey only break up the scat, but also so particles could 2015). Cookson, OK, is located in the eastern QRWVWD\GU\E\ÀRDWLQJRQWKHZDWHU¶VVXUIDFH portion of the state and is 16 km from where We then washed the samples through a series these 3 counties intersect. Our study site was of sieves (0.4 mm and 1 mm, H & C Sieving predominantly an oak-hickory forest with some systems, Models 6998 and 7003, Columbia, pine associations (Duck and Fletcher 1943). Maryland, USA) to separate particles to equal In addition to the public wildlife management VL]H DQG UHPRYH DQ\ XQZDQWHG PDWUL[ )RRG areas, much of the private land is used for LWHPV ZHUH LGHQWL¿HG PDFURVFRSLFDOO\ DV ZHOO wildlife management, though some landowners DVZLWKDGLVVHFWLQJPLFURVFRSH7KHLGHQWL¿HG raise livestock as well (Sara Lyda [Oklahoma LWHPV ZHUH WKHQ SODFHG LQ ¿OP FRQWDLQHUV IRU State University, Stillwater, Oklahoma] personal VWRUDJH:HFDWHJRUL]HGWKHLWHPVLQWRIRUDJH communication [August 2015]). groups: anthropogenic food (corn), reproductive SODQW SDUWV IUXLW VHHGV DQG ÀRZHUV  KHUEDJH As part of the larger study, bears were lured (roots, stems, and leaves), animal matter (bones, into barrel traps with donuts and other baked hair, and insect parts), debris (rocks, wood, bark, goods. Corn was also set outside of traps. Traps DQGSLQHQHHGOHV DQGXQLGHQWL¿DEOHPDWWHU were checked daily unless the forecast called for WHPSHUDWXUHVKLJKHUWKDQÛ&LQZKLFKFDVH :H LGHQWL¿HG DOO PDWWHU WR VSHFLHV ZKHQ the doors were removed from the traps. SRVVLEOH,QVHFWVZHUHLGHQWL¿HGWRFODVVGXHWR time constraints. To identify all plant material, For scat analysis we followed the same we used Field Guide to Oklahoma Plants (Tyrl PHWKRGVDV*UHHQOHDIHWDO  DQG*UDEHU et al. 2002) and the United States Department (1981). From May through November 2014 RI $JULFXOWXUH 3ODQW 'DWDEDVH 86'$ 15&6 we searched for scat near roads, trap sites, hair 2015). Many of the dietary items, such as a snares, natural forage areas, and potential trap majority of the grass specimens, were too Proc. Okla. Acad. Sci. 98: pp 37 - 45 (2018) 40 J.P. Connor, V.L. Jackson, J.R. Mittelhauser, and W.S. Fairbanks degraded to identify further than . The test (Zar 2009). This test was used for further majority of seeds were keyed to species. Bone analysis after running an ANOVA or Kruskal- IUDJPHQWVZHUHLGHQWL¿HGWRVSHFLHVJHQXVRU Wallis test and it was used to determine which family. The other fragments were labeled as JURXSVLQWKHVDPSOHZHUHVLJQL¿FDQWO\GL൵HUHQW large animal bones or unknown bones. We then The alpha level for tests was set at 0.05. measured percent volume and percent frequency of occurrence of each food item in scat samples 5HVXOWV WKDWZHUH!RIWKHYROXPHRIDVDPSOH *UDEHU *UDEHUDQG:KLWH*UHHQOHDIHWDO The majority of scat samples were found 2009). We measured percent volume of the 6 in Cherokee and Sequoyah counties with only classes using water displacement to the nearest 1 whole sample collected in Adair County. 1%. Volumetric analysis tends to overestimate Six samples were found in or near bear traps, the amount of herbage and underestimate easily 5 samples were found near blackberry (Rubus digested items such as reproductive plant parts spp.) patches, and 11 samples were collected like blackberries and animal tissue (Hatler near deer feeders. An estimated 306 hours were  0HDOH\  *UDEHU  7R DWWHPSW spent searching for scat samples. to more accurately assess dietary habits, we also calculated the percent frequency of occurrence From 16 May through 17 October 2014, a of food items as the percent of total scat samples total of 38 whole samples and 2 partial samples in which an item composed at least 1% of the were collected from 24 sites (8 samples in May, YROXPHRIDVDPSOH *UDEHU*UDEHUDQG 8 samples in June, 16 samples in July, 5 samples :KLWH*UHHQOHDIHWDO  in August, and 1 sample in October). The volume of each sample after being washed through the All scat that resembled bear scat was sieves ranged from 0.9 mL to 117 mL with an collected. Bear scat is variable in shape and is average volume of 35.3 mL per sample. There based on what the bear has eaten. If the animal ZHUHGL൵HUHQWGLHWDU\LWHPVFROOHFWHGIURPWKH has eaten more berries, the scat is more globular scat. There were 12 samples of the 38 collected and not as solid. If the scat is more herbage or that could have been small bear or based animal matter, the sample is more tubular and RQVL]HVKDSHDQGFRQWHQWV:HXVHGWKH0DQQ VROLG 6L]H LV DOVR YDULDEOH GHSHQGLQJ RQ WKH Whitney U test to compare these scat samples VL]HRIWKHDQLPDO&R\RWHVDUHRPQLYRURXVDQG to the rest of the samples that resembled only eat many of the same kinds of soft mast such bear, based on the volumetric amounts of each as blackberries, black cherries, and persimmons of the 6 food categories. These samples were (McVey et al. 2013), and their scat can look IRXQGWREHVLJQL¿FDQWO\GL൵HUHQWDQGWKH\ZHUH similar. Bear scat is more variable in shape, but excluded from further analyses (Mann Whitney tubular scats are common. Coyote scat is tubular 8WHVW3   DQGRIDVL]HVLPLODUWRDVPDOOEHDU7KLVLVD potential source of error. Items in the anthropogenic category were the most abundant item in terms of volume (Table The Mann-Whitney U test, a non-parametric 1). However, anthropogenic food items ranked 2-tailed test (Zar 2009), was used to compare second in frequency of occurrence in scat the volume of anthropogenic food items to the samples collected. The only anthropogenic item volume of natural food items. We also used the LGHQWL¿HGZDVFRUQ Zea mays). Mann-Whitney U test to compare food amounts from possible coyote scat against known bear The next most abundant food category VFDWV8VLQJ6366  ZHDOVRFRPSDUHGWKH was the reproductive plant parts (Table 1). volumetric amounts of the 6 diet groups against Reproductive plant parts were found most each other using the Kruskal-Wallis One-way frequently in samples collected. Blackberry $QDO\VLV RI 9DULDQFH LQFRUSRUDWLQJ 7XNH\¶V seeds, black cherry seeds (Prunus serotina), +RQHVW 6LJQL¿FDQW 'L൵HUHQFH +6'  SRVW KRF wild rye (Elymus VSS  LQÀRUHVFHQFH ZHUH WKH Proc. Okla. Acad. Sci. 98: pp 37 - 45 (2018) Black Bear Diet in Oklahoma 41 most abundant. frequency of deer feeders on the landscape, the abundance of corn in scat may be biased by the The majority of animal matter found was fact that corn was used to attract bears to trap insect parts, primarily ant and bee exoskeletons. sites, and some samples were collected near trap White-tailed deer (Odocoileus virginianus) hair sites. However, bears would have to be at the was found and Leporid and Microtus bones were trap site long enough for corn ingested at the also present, but in small amounts. trap site to pass through the digestive system for this to bias our results. Herbage was not very abundant in terms of volume but was abundant in terms of frequency. Reproductive plant parts were also very Blackberry leaves were the most abundant in abundant due to the fact that blackberries and this category. The next most abundant category black cherries begin producing mast throughout ZDV XQLGHQWL¿HG JUDVVHV 6RPH RI WKH JUDVVHV April-May (Tyrl et al. 2002). These plants are LGHQWL¿HG ZHUH SDQLF JUDVV Panicum spp.), common and attractive to wildlife. Seeds and blue stem (Andropogon spp.), and wheatgrass berries are common in the diet of brown bears (Agropyron spp.). (Ursus arctos  DQG EODFN EHDUV *UDEHU DQG White 1983). In Washington, berries are a major Debris was present in most scat samples. This SDUW RI WKH EHDUV¶ GLHW /LQG]H\ DQG 0HVORZ category consisted of small rocks, bark, wood 1977). FKLSV DQG SLQH QHHGOHV 8QLGHQWL¿DEOH PDWWHU made up a very small portion and consisted Animal matter was somewhat common. of anything that was too decomposed to be Though the amount of animal matter by volume LGHQWL¿HG was not very large, animal remains were frequently found. The majority of this was insect The volume of anthropogenic food in scats parts. Bears frequently eat ants (Chatelain 1950; GLGQRWGL൵HUIURPWKHYROXPHRIQDWXUDOIRRG +DWOHU%HHPDQDQG3HOWRQ DQGZDVSV 0DQQ:KLWQH\ 8 WHVW 3   1   %\ +DWOHU%R\HU%HHPDQDQG3HOWRQ comparing median ranks by ranking food   7KH PDMRULW\ RI LQVHFWV LGHQWL¿HG ZHUH categories by volume in all samples, the 6 diet ants but there were also bee and wasp remains. JURXSV ZHUH VLJQL¿FDQWO\ GL൵HUHQW .UXVNDO Bears could have been seeking out ants or they :DOOLV 3    DQG ZHUH VHSDUDWHG LQWR  could have ingested the ants with food items statistically distinct groups: Anthropogenic food picked up from the ground. White-tailed deer hair > Reproductive plant parts > Animal matter ZDVQ¶WYHU\SUHYDOHQW YROXPHRUIUHTXHQF\ LQ  +HUEDJH  'HEULV  8QLGHQWL¿HG 7XNH\¶V the scat samples. The seasonal abundance of soft +RQHVW6LJQL¿FDQW'L൵HUHQFHWHVWDOSKD   mast in the environment could attract bears more so than cervid remains, making animal tissue 'LVFXVVLRQ less prevalent in scat samples. Cervid remains in black bear scat usually represent carrion and 7KH DQWKURSRJHQLF IRRG LGHQWL¿HG LQ WKH MXYHQLOHV 6FKZDUW] DQG 0LWFKHOO  7LVFK scat from bears in our study was corn most 1961; Hatler 1972), but black bears are capable likely obtained from deer feeders rather than of preying on adult white-tailed deer (Svoboda cropland. Many of the landowners manage and/ et al. 2011). or hunt deer over bait, and they do so with deer IHHGHUV %HDUKXPDQ FRQÀLFW DULVHV ZKHQ WKH Herbage was not a major component in bears destroy feeders or break into the feeders. volume and frequency in the samples. We Landowners try to deter bears from breaking believe this is because of the time of the year the into the deer feeders using several methods VFDWVZHUHFROOHFWHG*UDEHUDQG:KLWH   VXFK DV UDLVLQJ IHHGHUV IDU R൵ WKH JURXQG E\ determined the amount of herbage decreases cables. Many times bears are still able to breach from spring to fall. Herbage is a very important the container and obtain the corn. Despite the dietary item in the spring, and it can make up Proc. Okla. Acad. Sci. 98: pp 37 - 45 (2018) 42 J.P. Connor, V.L. Jackson, J.R. Mittelhauser, and W.S. Fairbanks 7DEOH&RQWHQWVRI$PHULFDQEODFNEHDUVFDWFROOHFWHGLQ$GDLU&KHURNHHDQG6HTXR\DK FRXQWLHV0D\WKURXJK1RYHPEHU1  )RRG5HVRXUFH 9ROXPH )UHTXHQF\

Anthropogenic 54.4% 61.2%

Corn 54.4% 61.2%

5HSURGXFWLYH3ODQW3DUWV 32.3% 69.2%

Blackberry 26.2% 34.6%

Black Cherry 1.6% 15.4%

Wild Rye 3.7% 19.2%

$QLPDO0DWWHU 4.2% 69.2%

Insect 2.4% 61.5%

Deer hair 1.2% 11.5%

+HUEDJH 2.4% 46.2%

Blackberry leaves 0.9% 15.4%

*UDVV 0.4% 15.4%

Bluestem grass 0.4% 3.9%

'HEULV 6% 80.8%

8QLGHQWLILDEOH 0.8% 15.4%

 DERXW KDOI RI WKHLU GLHW 3LHNLHOHN DQG %XUWRQ GRHV QRWD൵HFW*577KHUHZHUH GD\V ZKHQ %R\HU(DJOH 2XU¿UVWVDPSOH multiple samples were collected from a single was not collected until May, so herbage may be site. Although cameras often indicated more underrepresented because the ripening of some than one bear visiting the trap sites in a 24-h berry species may have already attracted the period, our samples should not be considered bears away from herbage. independent.

This study was limited in terms of scat Similar diet studies have been conducted collection and therefore was not a complete LQ

This could mean bears are searching out corn Benson JF and Chamberlain MJ. 2006. Food over natural food sources or that deer feeders habits of Louisiana black bears (Ursus are abundant and bears use them as encountered. americanus luteolus) in 2 subpopulations of If bears are searching for deer feeders, this may the Tensas River Basin. American Midland OHDGWRFRQÀLFWZKLFKFDQFRQWULEXWHWRFRQÀLFW Naturalist 156:118–127. between humans and bears. Boyer KB.1976. Food habits of black bears (Ursus americanus) in the Banning Canyon This study showed that bear diet in the area of San Bernardino National Forest. 2NODKRPD 2]DUNV DUHD ZDV D൵HFWHG E\ WKH 06 7KHVLV &DOLIRUQLD 6WDWH 3RO\WHFKQLF availability of anthropogenic food resources. 8QLYHUVLW\3RPRQDSS The bear population in this area is expanding Breck SW, Lance N, and Seher V. 2009. Selective into a human-dominated landscape, which foraging for anthropogenic resources by black LQFUHDVHV WKH OLNHOLKRRG RI FRQÀLFW  :LWK WKLV EHDUVPLQLYDQVLQ

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Proc. Okla. Acad. Sci. 98: pp 37 - 45 (2018) Black Bear Diet in Oklahoma 45

6LNHV 56 *DQQRQ :/ DQG WKH$QLPDO &DUH 7\UO 5- %LGZHOO 7* DQG 0DVWHUV 5(  and Use Committee of the American Society Field guide to Oklahoma plants, commonly RI 0DPPDORJLVWV  *XLGHOLQHV RI WKH encountered prairie, shrubland, and forest American Society of Mammalogists for the Species. Oklahoma State University. Stillwell, use of wild mammals in research. J Mammal Oklahoma. 92: 235–253. U.S. Census Bureau. 2016. Human population 6PLWK .* DQG &ODUN -'  %ODFN EHDUV for counties in Oklahoma [online]. Available in Arkansas: characteristics of a successful from: Census.gov. (Accessed January 26, translocation. J Mammal 75:309-320. 2018). 6366,QF5HOHDVHG3$6:6WDWLVWLFVIRU USDA [United States Department of :LQGRZV9HUVLRQ&KLFDJR6366,QF Agriculture], NRCS [Natural Resources Svoboda NJ, Belant JL, Beyer DE, Duquette &RQVHUYDWLRQ 6HUYLFH@  7KH 3/$176 JF, Stricker HK, and Albright CA. 2011. Database. Available from: http://plants.usda. predation of an adult gov. (Accessed 6 October 2015). white-tailed deer. Ursus 22:91-94. :RRGUR൵H53UHGDWRUVDQGSHRSOH8VLQJ Swenson JE, Sandegreen F, and Soderberg human densities to interpret declines of large $  *HRJUDSKLF H[SDQVLRQ RI DQ carnivores. Animal Conserv 3: 165–173. increasing population: evidence =DU -+  %LRVWDWLVWLFDO DQDO\VLV 3UHQWLFH for presaturation dispersal. J of Animal Ecol Hall, New Jersey 663 pp. 67:819-826. Submitted September 10, 2018 Accepted November 6, 2018 Tisch EL. 1961. Seasonal food habits of WKH EODFN EHDU LQ WKH :KLWH¿VK 5DQJH RI northwestern Montana. M.S. Thesis. Montana State University, Missoula. 108pp.

Proc. Okla. Acad. Sci. 98: pp 37 - 45 (2018)