Regular issues
Monographs
Special Issues



Southeastern Naturalist
    SENA Home
    Range and Scope
    Board of Editors
    Staff
    Editorial Workflow
    Publication Charges
    Subscriptions

Other EH Journals
    Northeastern Naturalist
    Caribbean Naturalist
    Neotropical Naturalist
    Urban Naturalist
    Eastern Paleontologist
    Journal of the North Atlantic
    Eastern Biologist

EH Natural History Home

Seasonal Diets of an Introduced Population of Fallow Deer on Little St. Simons Island, Georgia
Brian W. Morse, Mandi L. McElroy, and Karl V. Miller

Southeastern Naturalist, Volume 8, Number 4 (2009): 571–586

Full-text pdf (Accessible only to subscribers.To subscribe click here.)

 

Site by Bennett Web & Design Co.
2009 SOUTHEASTERN NATURALIST 8(4):571–586 Seasonal Diets of an Introduced Population of Fallow Deer on Little St. Simons Island, Georgia Brian W. Morse1, Mandi L. McElroy1, and Karl V. Miller1,* Abstract - We examined seasonal diets of an introduced population of Dama dama dama (European Fallow Deer) on Little St. Simons Island, GA. We analyzed rumen contents from hunter-harvested deer during fall and winter of 2004–05 and 2005–06. Fecal pellets also were collected monthly from November 2004 to December 2005 and examined microscopically for unique plant-cell characteristics. Fallow Deer utilized a variety of food items based on seasonal availability, although mast and browse were the most abundant food items in the rumens examined. Fallow Deer preferred Quercus spp. (oak) acorns, but consumed more Sabal palmetto (Cabbage Palm) fruit when acorns were less abundant. Microhistological techniques underestimated the occurrence of highly digestible items such as mast, but were more effective at identifying grasses and browse. Grasses were the most common and abundant forage class in feces, with peak grass use in the summer (67%). Fallow Deer’s ability to utilize a wide variety of food items including low-quality forage has contributed to their success in this barrier island ecosystem. However, low productivity, suppressed body weights, and small antler characteristics are likely due to a low-quality diet and over-population. Introduction Dama dama dama L. (European Fallow Deer) is one of the most widely distributed cervid species in the world. Conjectured to have come originally from southeastern Europe and adjacent Asia Minor, this subspecies of Fallow Deer has been the subject of repeated introductions since ancient times (Chapman and Chapman 1975). Today it inhabits Europe, Asia, North America, South America, Africa, and Australia (Chapman and Chapman 1975). In the United States, Fallow Deer have been released in Georgia as well as Alabama, California, Colorado, Florida, Kentucky, Louisiana, Massachusetts, Mississippi, Nebraska, New Mexico, New York, Oregon, Texas, Utah, and Virginia (Chapman and Chapman 1975, Mungall 2000). Only a few free-ranging populations remain. One of these exists on Little St. Simons Island (LSSI), a coastal barrier island near Brunswick, GA. The potential impacts of exotic deer populations on native deer species and local ecosystems are a growing concern (Davidson et al. 1985, Jackley 1991, Keiper 1985, USDOI NPS 2006). Recently, the National Park Service (NPS) has recommended removal of exotic Fallow Deer and Axis axis Erxleben (Axis Deer) from Point Reyes National Seashore, California (USDOI NPS 2006). Contributing factors in this decision stemmed from declines in native Odocoileus hemionus columbianus Richardson (Black-tailed Deer) 1D.B. Warnell School of Forestry and Natural Resources, University of Georgia, Athens, GA 30602. *Corresponding author - kmiller@warnell.uga.edu. 572 Southeastern Naturalist Vol. 8, No. 4 and Cervus elaphus nannodes Merriam (Tule Elk) populations, as well as habitat degradation within the Park (USDOI NPS 2006). On LSSI, there is concern that Fallow Deer are inhibiting the regeneration of maritime forest and threatening other sensitive barrier island ecosystems. Fallow Deer were first introduced to LSSI during the 1920s (S. Johnson, Bronx Zoo, New York, NY, pers. comm.). In the early 1980s, in an effort to reduce damage to native ecosystems, cattle were removed from the island and a hunting program was initiated to reduce Fallow Deer numbers. Current Fallow Deer density on LSSI is approximately 47 deer/km2 (B.W. Morse, unpubl. data), and hunting is the primary cause of mortality in the population. The historical status of native Odocoileus virginianus Zimmermann (White-tailed Deer) is unknown. Sightings of White-tailed Deer on LSSI in the recent past have been rare, and none were observed from 2002–2006. However, White-tailed Deer are present on all the surrounding barrier islands as well as mainland Georgia, and are likely to have been displaced from LSSI by the exotic Fallow Deer. Little is known about the ecology of Fallow Deer on LSSI, including their food habits and the potential negative effects browsing may have on the barrier island ecosystem. Our objective was to document seasonal food habits of Fallow Deer on LSSI. An understanding of food habits is important when managing an exotic herbivore, especially on an island where resources are limited. If suboptimal foods comprise the majority of the diet, it is likely that higher-quality food items are over-exploited by an overabundant population (Bruno and Apollonio 1991). Study Area Little St. Simons Island is a privately owned, 4340-ha coastal barrier island in Glynn County, GA approximately 21 km northeast of Brunswick. The island is bordered by the Atlantic Ocean to the east, the Altamaha River to the north, and the Hampton River to the south and west, with no land linkage. The island is typical of other undeveloped barrier islands in the South Atlantic/Carolinian biogeographic classification (McIntyre 2001). Most of the island (3189 ha) consists of tidally infl uenced salt marshes dominated by Spartina alternafl ora Loisel. (Smooth Cordgrass) in lower marshes and Distichlis spicata L. (Saltgrass) in higher marshes. Upland habitats include primary and secondary dune systems, maritime shrub communities dominated by Morella cerifera (L.) Small (Wax Myrtle), mixed pine-oak-palmetto forest, and mature maritime forests of Quercus virginiana P. Mill. (Live Oak) and Q. laurifolia Michx. (Laurel Oak). Freshwater habitats are limited to a few permanent ponds and ephemeral sloughs. Floral diversity is low compared to nearby larger islands and the mainland because of the island's small size, poor soils (Rigdon and Green 1980), and topographic limitations. Browsing pressure by Fallow Deer is another potential cause. Elevations range from sea level to 9 m above mean sea level (MSL); however, most of the upland habitats are less than 3 MSL. Marine infl uences provide 2009 B.W. Morse, M.L. McElroy, and K.V. Miller 573 cooler summers and warmer winters than the mainland (McIntyre 2001). January and July temperatures average 11.9 ºC and 28.1 ºC, respectively, and mean annual rainfall is 132.4 cm (Southeast Regional Climate Center 2007). Severe droughts and fl oods are common. Flooding occurs mainly from tropical storm systems and high spring tides. Fallow Deer are the only large free-ranging mammal on LSSI (except for rare sightings of White-tailed Deer). Predatory mammals such as Lynx rufus Schreber (Bobcat), Vulpes vulpes L. (Red Fox), Urocyon cineroargenteus Schreber (Gray Fox), and Canis latrans Say (Coyote) are absent. The only potential predator of Fallow Deer is Alligator mississippiensis Daudin (American Alligator). Competition for mast resources includes Sciurus carolinensis Gmelin (Gray Squirrel) and Procyon lotor L. (Raccoon). Methods We assessed Fallow Deer food habits by macroscopic rumen analysis, microhistological analysis of fecal pellets, and direct observation. We sampled rumen contents from hunter-harvested deer on LSSI from October 2004 to February 2005 (Year 1) and October 2005 through mid-March 2006 (Year 2). Individual rumen samples were separated into fall and winter seasons. The fall season consisted of deer harvested in October, November, or December; winter consisted of deer harvested in January, February, and March. We examined the contents of 54 rumens: 18 in fall 2004, 18 in winter 2005, 7 in fall 2005, and 11 in winter 2006. Rumen analysis followed Puglisi et al. (1978). Immediately after harvest, rumens were opened, and their contents thoroughly mixed. One-liter samples were taken from each rumen, frozen in plastic containers, and labeled appropriately. Later, rumen samples were thawed and washed with water through 2 stacked sieves (mesh sizes of 3.36 mm and 5.66 mm). Food items were separated according to plant type and species. Once separated, each plant was identified by eye, hand lens, or dissecting microscope and checked against a reference collection prepared by gathering plant specimens from LSSI. We identified each plant item to the lowest possible taxon. Unidentifi- able fragments were categorized as unknown. We determined the volume of each food by water displacement. We calculated percent seasonal occurrence by dividing frequency of occurrence of each food item (or group) by the number of rumen samples for each season. Volumes were determined using the aggregate volume method described by Martin et al. (1946). Aggregate volume is calculated by summing the volume of a particular food item in all rumen samples and dividing by the total volume of all foods in all samples, giving importance to absolute volume of food consumed by all individuals in the sample (Litvaitis et al. 1996). We considered food items found in rumens to be important if they constituted greater than 5% of the aggregate volume, as well as greater than 50% frequency of occurrence. Because we could not collect rumen samples during spring and summer, we collected fecal pellets every month from November 2004 to December 574 Southeastern Naturalist Vol. 8, No. 4 2005 to examine spring and summer diet, as well as to compare fall and winter diets using the 2 techniques. To obtain a representative sample from all parts of the island, we divided the island into 6 sections based on geography, and collected one pellet group per month from each section. We evaluated pellets in the field for relative moisture content and decomposition to ensure that samples were deposited within the sample month. Each pellet group represented one deer. Pellets were placed in plastic bags with iodized salt (J. Rentfl eish, Micro Composition Laboratory, Ft. Collins, CO, pers. comm.), labeled, and shipped to Micro Composition Laboratory for analysis. Fecal contents were analyzed using microhistological techniques described in detail by Johnson et al. (1983). Pellets were ground in a Wiley mill, and one slide was prepared. Twenty fields were read for each sample using a 100x binocular microscope. Plant fragments were identified based on epidermal tissue characteristics using laboratory reference slides and a plant list from LSSI. All identifiable plant fragments in the 20 fields of view were counted for each pellet group. We calculated relative percent density (number of fragments of a plant divided by the number of all fragments in all samples) for each pellet group. Items from rumen and fecal samples that could not be placed in fruit, grass, browse, or forb categories were combined in an “other” category. Months were grouped into seasons for comparison. Fall and winter months are as listed above, with spring season being April, May, and June, and summer being July, August, and September. Food habits data collection methods were approved by the University of Georgia Institutional Animal Care and Use Committee (Permit # A2005-10150-0). Results We were able to categorize >88% of the food items recovered from rumens to the familial or lower taxonomic level. Vitis spp. (Grapes) were the only food found in 2004 that did not occur in 2005. However, we recovered 10 foods in 2005 that did not occur in 2004. Fruit constituted the greatest volume in rumen samples except during winter 2005. Fruit use was greater in Year 2 than Year 1, and was greater than the combination of all other food groups in fall of 2005 and winter of 2006 (Fig. 1). Quercus spp. acorns occurred in >55% of Fallow Deer rumens in both seasons and years, and >72% in winter 2005 and Year 2. Acorns were found in 100% of fall 2005 rumens, and represented 97% of the volume of all fruit consumed. The frequency of use and total volume consumed was greater in 2005 than 2004. The only other fruits that represented >5% in any season or year were those of Sabal palmetto (Walt.) Lodd. ex J.A. & J. H. Schultes (Cabbage Palm) and Prunus caroliniana (P. Mill.) Ait. (Carolina Laurelcherry) (Appendix1). Browse was the second most abundant forage class in Fallow Deer rumens in fall 2005 and winter 2006 (Fig. 1). Seventeen browse items were detected. Browse items tended to be a more important component of the diet in Year 1 2009 B.W. Morse, M.L. McElroy, and K.V. Miller 575 than Year 2, both in terms of frequency of occurrence and in volume. During both years, Wax Myrtle, Ilex vomitoria Ait. (Yaupon), and Live Oak leaves were the most common browse items. Grape foliage and Phoradendron serotinum (Raf.) Reveal & M.C. Johnston (Mistletoe) were important components of the diet in 2004, but not in 2005. Various forbs, primarily Pleopeltis polypodioides (L.) Andrews & Windham (Resurrection Fern), Tillandsia usneoides L. (Spanish Moss), and Polygonum sp. (smartweed) were common in Fallow Deer rumens in all seasons, but rarely comprised a significant portion of the diet (Appendix 1). Grasses, sedges, and rushes were commonly recovered from deer rumens, but only constituted a major component of the diet during fall (Fig. 1). Likewise, fungi and lichens (in “other” category) were a common, but relatively unimportant component of the diet. Microhistologic analysis of fecal pellets revealed 72 foods consumed (Appendix 2). Grasses were consumed during all seasons, and were consumed more frequently during spring and summer than other forage classes (Fig. 2). Grass occurred in 67% of summer diets of Fallow Deer on LSSI. Uniola spp. (seaoats) was the most commonly observed grass in fecal samples. Cynodon dactylon (L.) Pers. (Bermudagrass), Cyperus spp. (sedges), Salt Grass, and Juncus sp. (rush) also were common forage items revealed from fecal samples. Eragrostis sp. (Lovegrass) was an important food item in fall 2004 (9% RD). Leaves and stems from 20 woody plant species were identified in the fecal samples. Oak leaves, most probably Live Oak, Laurel Oak, or Q. geminaa Small (Sand Live Oak), were the most common browse item, typically comprising half of the relative percent density of all browse items recovered Figure 1. Aggregate percent volume and 95% CI of forage classes in rumens of Fallow Deer from Little St. Simons Island, GA, 2004–2006. 576 Southeastern Naturalist Vol. 8, No. 4 in feces. Juniperus virginiana L. (Eastern Redcedar) and Salix sp. (willow) were eaten in all seasons (Appendix 2). Twenty-two species of forbs were detected in the feces of Fallow Deer (Appendix 2). Only Croton punctatus Jacq. (Gulf Croton) and an unknown forb species occurred year-round. Gulf Croton was an important food item in winter. Another unknown forb species contributed >5% of the diet in spring and summer seasons. Other items identified in Fallow Deer feces included a corn kernel, an unknown fl ower, a grass seed and glume, a legume pod, an unknown seed, and lichen. Our observational data provided additional insights into Fallow Deer diets on LSSI. In May 2004, we observed 3 bucks foraging exclusively on fallen Magnolia grandifl ora L. (Southern Magnolia) fl ower petals. In August 2004, a buck readily consumed a locally abundant white mushroom. In August 2005, we observed a buck feeding in a salt marsh on Smooth Cordgrass. An observation of a yearling buck feeding on fallen Sideroxylon tenax L. (Tough Bumelia) fruits occurred in November 2005. Additionally, we observed obvious browse or browse lines on Smilax spp. (greenbriar), Live oak, Carolina Laurelcherry, Eastern Redcedar, Erythrina herbacea L. (Eastern Coralbean), and Forestiera segregata (Jacq.) Krug & Urban (Florida Swampprivet). Seeds of Amaranthus sp. (pigweed) were collected from the abomasum of one deer in December 2006. Discussion Food habits of Fallow Deer have been reported from various localities, and results differ among study areas. Graminoids compose the majority of Fallow Deer diets in California (Wehausen and Elliott 1982), Poland (Borkowski and Obidziński 2003), and Spain (Garcia-Gonzalez and Cuartas Figure 2. Mean percent relative density and 95% CI of forage classes by season from feces of Fallow Deer on Little St. Simons Island, GA, 2004–2005. 2009 B.W. Morse, M.L. McElroy, and K.V. Miller 577 1992). In contrast, browse was reported as the dominant forage in New Zealand (Nugent 1990), Italy (Bruno and Apollonio 1991), and Texas (Butts et al. 1982). On LSSI, Fallow Deer consumed a variety of food items based on seasonal and annual availability. Our results also varied according to analytical technique used. Techniques for examining herbivore diets have been thoroughly reviewed by Holechek et al. (1982). Fecal analysis typically is more effective for species identification, whereas rumen analysis provides an assessment of the relative importance of common food items, as well as a more accurate estimate of consumption of highly digestible food items such as fruits and fungi. Based on rumen analysis, fruit was important forage during fall and winter. The increase in fruit use from 2004 to 2005 refl ects the abundant mast crop observed in 2005 (Fig. 3). During 2005, mast, primarily fallen acorns, comprised more than half of the rumen content volume in both fall and winter, similar to reports of acorn use by Fallow Deer in England (Chapman and Chapman 1975). Although acorns are a highly preferred food item, mast from Cabbage Palms is consumed frequently when acorns are less abundant. Cabbage Palm fruits represented >10% of the fall and winter diets in 2004, but use dropped to less than 1% in 2005 when acorns were anecdotally determined to be more abundant (Fig. 3). Similar use of Cabbage Palm fruits has been reported for White-tailed Deer on nearby barrier islands when acorns were less abundant (Osborne et al. 1992, Warren et al. 1990). Mast failures on these barrier islands can have dramatic impacts on White-tailed Deer fitness, reproduction, movements, and survival (Osborne et al. 1992). Considering the heavy use found in the present study, mast failures on barrier islands may cause similar effects in Fallow Deer. Figure 3. Aggregate percent volume and 95% CI for Quercus spp. and Sabal palmetto fruits from rumens of Fallow Deer on Little St. Simons Islands, Georgia 2004–2006. 578 Southeastern Naturalist Vol. 8, No. 4 Fruiting bodies were not revealed by microscopic fecal pellet examination because fruits are generally succulent, palatable, and highly digestible in ruminant digestive tracts (Anthony and Smith 1974). Food items remaining in feces are less digestible and provide bias towards these items. Thus, subtotals of each forage class found in the feces should probably be viewed as an index of non-mast food items consumed by deer. The amount of browse in rumens declined from 2004 to 2005, refl ecting the increase in acorn mast consumption. Osborne et al. (1992) reported White-tailed Deer used Wax Myrtle when other foods were exhausted, indicating low preference. Although Fallow Deer preference of Wax Myrtle is unknown, its abundance on LSSI and high frequency of occurrence in rumen samples indicate that it is an important food item, especially when other foods are limited. Unexpectedly, Wax Myrtle was not detected in the fecal samples, perhaps refl ecting an error in microhistological identification of this species. There is concern that Fallow Deer on LSSI are inhibiting Live Oak regeneration in the maritime forest (W. Paulson, owner of LSSI, pers. comm.), a community dominated by Live Oak and Laurel Oak. Both rumen and fecal pellet analysis indicated use of Live Oak in all seasons with peaks in winter. Browse lines observed on the island provide additional evidence of Live Oak use by Fallow Deer. Browse quality generally decreases as plants mature, and deer typically select younger, more digestible plants and plant parts (Cypher and Cypher 1988). By selecting young Live Oak seedlings, Fallow Deer certainly could impact regeneration of this species. Oak species, including Live Oak, were considered important foods in the winter diets of Fallow Deer in Texas (Jackley 1991). Browse lines and absence of regeneration can indicate population overabundance and overuse of food resources (Anonymous 1994, Healy 1997). Browse lines and browsing were evident in certain vegetative communities of LSSI. Browse lines on Florida Swampprivet is also of particular concern. This shrub is listed as rare (S2) in Georgia (Chafin 2007), and should be monitored for browsing impacts and protected if necessary. Forbs were frequently used, but contributed little to the diet compared to other food items. In California, forbs are an important component of the diet of introduced Fallow Deer (Wehausen and Elliot 1982), whereas a Texas study where other ruminants were excluded indicated forbs contributed 12% of the diet (Butts et al. 1982), similar to our results. Based on fecal analyses, we did not observe dramatic seasonal fl uctuations in forb use as has been reported in other studies (Garcia-Gonzalez and Cuartas 1992, Warren et al. 1990). Succulent forbs are highly digestible, making identification difficult and causing an underestimation in abundance (Litvaitis et al. 1999). Fallow Deer show a marked preference for graminoids across their range (Garcia-Gonzalez and Cuartas 1992, Jackson 1977, Kerridge and Bullock 1991, Putman et al. 1993). The occurrence and volume of grass in Fallow Deer rumens declined between fall and winter in response to decreased 2009 B.W. Morse, M.L. McElroy, and K.V. Miller 579 availability and palatability of warm-season grasses. Fallow Deer may be more efficient than native White-tailed Deer in digesting fiber (Hofmann 1985), and have greater relative rumen capacity than White-tailed Deer, allowing Fallow Deer to utilize a grass-dominated diet more efficiently (Henke et al. 1988). Uniola sp. was the most important grass consumed in all seasons. The only species of Uniola currently known from LSSI is Uniola paniculata L. (Sea Oats), which has received protection by the Sea Oats Conservation Act of 1973 (O.C.G.A. 12-5-310 to 12-5-312). However, due to similarities in cellular characteristics with Chasmanthium laxum (L.) Yates (Slender Woodoats), we cannot differentiate between these species. Although Fallow Deer are not commonly seen in the fore-dune ridge areas where Sea Oats occur, future monitoring of Fallow Deer use of this species is warranted. Our results confirm the dietary adaptability of Fallow Deer and their ability to utilize a variety of food resources efficiently. Dietary preferences, coupled with high deer densities on LSSI (Morse and Miller 2009) may be altering the plant community composition and structure. Our data will be most useful in combination with vegetation surveys to examine forage availability on LSSI to assess the impact Fallow Deer have on vegetative communities and plant species of special interest. Acknowledgments We thank M. Pollock for assistance in field data collection, D. Osborn for support and advice, and the cooperation of Little St. Simons Island, LLC and their staff. M. Dodd of the Georgia Department of Natural Resources and K. Keel of the Southeastern Cooperative Wildlife Disease Study provided us with access to facilities for conducting analysis. P. Hale assisted with rumen-content plant identification. Funding was provided by the University of Georgia, the Bobolink Foundation, the Coastal Barrier Island Foundation, and the Lowcountry Chapter of Safari Club International. Literature Cited Anonymous. 1994.The use and management of browse in the Edwards Plateau of Texas. US Department of Agriculture, Natural Resources Conservation Service, Temple, TX. 7 pp. Anthony, R.G., and N.S. Smith. 1974. Comparison of rumen and fecal analysis to describe deer diets. Journal of Wildlife Management 38:535–540 Borkowski, J., and A. Obidziński. 2003. The composition of the autumn and winter diets in two Polish populations of Fallow Deer. Acta Theriologica 48:539–546. Bruno, E., and M. Apollonio. 1991. Seasonal variations in the diet of adult male fallow deer in a sub-Mediterranean coastal area. Revue D’Ecologie 46:349–362. Butts, G.L., M.J. Anderegg, W.E. Armstrong, D.E. Hormel, C.W. Ramsey, and S.H. Sorola. 1982. Food habits of five exotic ungulates on Kerr Wildlife Management Area, Texas. Texas Parks and Wildlife Department, Technical Series Number 30, Austin, TX. 47 pp. Chafin, L.G. 2007. Field Guide to the Rare Plants of Georgia. State Botanical Garden of Georgia, University of Georgia, Athens, GA. 526 pp. 580 Southeastern Naturalist Vol. 8, No. 4 Chapman, D., and N. Chapman. 1975. Fallow Deer: Their History, Distribution, and Biology. T. Dalton, Ltd., Lavenham and Suffolk, Great Britain, UK. 271 pp. Cypher, B., and A. Cypher. 1988. Ecology and management of White-tailed Deer in northeastern coastal habitats: A synthesis of the literature pertinent to national wildlife refuges from Maine to Virginia. US Fish and Wildlife Service Biological Report. 88. 52 pp. Davidson, W.R., J.M. Crum, J.L. Blue, D.W Sharp, and J.H. Phillips. 1985. Parasites, diseases, and health status of sympatric populations of Fallow Deer and Whitetailed Deer in Kentucky. Journal of Wildlife Disease 21:153–159. Garcia-Gonzalez, R., and P. Cuartas. 1992. Food habits of Capra pyrenaica, Cervus elaphus, and Dama dama in the Cazorla Sierra (Spain). Mammalia 56:195–202. Healy, W.M. 1997. Infl uence of deer on the structure and composition of oak forests in Central Massachusetts. Pp. 249–266, In W.J. McShea, H.B. Underwood, and J.H. Rappole (Eds.). The Science of Overadundance: Deer Ecology and Population Management. Smithsonian Institution Press, Washington, DC. 402 pp. Henke S.E., S. Demarais, and J.A. Pfister. 1988. Digestive capacity and diets of White-tailed Deer and exotic ruminants. Journal of Wildlife Management 52:595–598. Hofmann, R.R. 1985. Digestive physiology of the deer. Their morphophysiological specialization and adaptation. Bulletin of the Royal Society of New Zealand 26:481–501. Holechek, J.L., M. Vavra, and R.D. Pieper. 1982. Botanical composition determination of range herbivore diets: A review. Journal of Range Management 35:309–315. Jackley, J.J. 1991. Dietary overlap among Axis, Fallow, Sika, and White-tailed Deer in the Edwards Plateau region of Texas. M.Sc. Thesis. Texas Tech University, Lubbock, TX. 189 pp. Jackson, J. 1977. The annual diet of the Fallow Deer (Dama dama) in the New Forest, Hampshire, as determined by rumen-content analysis. Journal of Zoology, London 181:465–473. Johnson, M.K., H. Wofford, and H.A. Pearson. 1983. Microhistological techniques for food habits analysis. US Department of Agriculture, Forest Service, Southern Forest Experiment Station. Research Paper SO-199. 40 pp. Keiper, R.R. 1985. Are Sika Deer responsible for the decline of White-tailed Deer on Assateague Island, Maryland? Wildlife Society Bulletin 13:144–146. Kerridge, F.J., and D.J. Bullock. 1991. Diet and dietary quality of Red Deer and Fallow Deer in late summer. Journal of Zoology, London 224:333–337. Litvaitis, J.A., K. Titus, and E.M. Anderson. 1996. Measuring vertebrate use of terrestrial habitats and foods. Pp. 254–274, In T.A. Bookhout (Ed.). Research and Management Techniques for Wildlife and Habitats. The Wildlife Society, Bethesda, MD. 740 pp. Martin, A.C., R.H. Gensch, and C.P. Brown. 1946. Alternative methods in upland gamebird food analysis. Journal of Wildlife Management 10:8–12. McIntyre, K.M. 2001. Little Saint Simons Island ecological characterization and resource management plan. Prepared for Little Saint Simons Island, LLC. (Unpublished report). Morse, B.W., and K.V. Miller. 2009. Population characteristics of an insular Fallow Deer (Dama dama) population on Little St. Simons Island, GA, USA. Wildlife Biology in Practice 5(1):1–10. 2009 B.W. Morse, M.L. McElroy, and K.V. Miller 581 Mungall, E.C. 2000. Exotics. Pp. 736–764, In S. Demarais and P.R. Krausman (Eds.). Ecology and Management of Large Mammals in North America. Prentice Hall, Upper Saddle River, NJ. Nugent, G. 1990. Forage availability and the diet of Fallow Deer (Dama dama) in the Blue Mountains, Otago. New Zealand Journal of Ecology 13:83–95. Osborne, J.S., P.E. Hale, R.L. Marchinton, C.V. Vansant, and J.M. Wentworth. 1992. Population ecology of the Blackbeard Island White-tailed Deer. Bulletin 26. Tall Timbers Research Station, FL. 108 pp. Puglisi, M.J., S.A. Liscinsky, and R.F. Harlow. 1978. An improved methodology of rumen-content analysis for White-tailed Deer. Journal of Wildlife Management 42:397–403. Putman, R.J., S. Culpin, and S.J. Thirgood. 1993. Dietary differences between male and female Fallow Deer in sympatry and in allopatry. Journal of Zoology, London 229:267–275. Rigdon, T.A., and A.J. Green. 1980. Soil survey of Camden and Glynn counties, Georgia. US Department of Agriculture, Soil Conservation Service, Washington DC. 82 pp. Southeast Regional Climate Center (SERCC). 2007. Brunswick, GA historical climate summary 2003–2006. Available online at http://sercc.com/climateinfo/ historical/historical_ga.html. Accessed 15 October 2008. Warren, R.J., S.K. Miller, R.D. Rowland, C.L. Rogers, and N.M. Gobris. 1990. Population ecology of White-tailed Deer on Cumberland Island National Seashore. Final Report. US National Park Service, Southeast Region, Atlanta, GA. 120 pp. Wehausen, J.D., and H.W. Elliott. 1982. Range relationships and demography of Fallow and Axis Deer on Point Reyes National Seashore. California Fish and Game 68:132–145. US Department of the Interior, National Park Service (USDOI NPS). 2006. Point Reyes National Seashore non-native deer management plan, Final environmental impact statement. Point Reyes National Seashore, Point Reyes, CA. 431 pp. 582 Southeastern Naturalist Vol. 8, No. 4 Appendix 1.Fall and winter rumen contents (percent occurrence [% Occ.] and percent volume [Vol.]) of Fallow Deer on Little St. Simons Island, GA, 2004–2006. Oct–Dec 2004 Jan–Feb 2005 Oct–Dec 2005 Jan–Mar 2006 n = 18 n = 18 n = 7 n = 11 Food item (species) Common name % Occ. Vol. % Occ. Vol. % Occ. Vol. % Occ. Vol. Fruit Quercus spp. Oak 55.6 8.0 83.3 25.4 100.0 55.9 72.7 30.4 Phoradendron serotinum (Raf.) Reveal & M.C. Johnston Mistletoe 9.1 0.1 Prunus carolinana (P. Mill.) Ait. Cherrylaurel 14.3 0.4 27.3 30.0 Serenoa repens (Bartr.) Small Saw Palmetto 14.3 0.2 9.1 0.9 Ilex vomitoria Ait. Yaupon Holly 14.3 0.6 18.2 1.0 Sabal palmetto (Walt.) Lodd. Ex J.A. & J.H. Schultes Cabbage Palm 72.2 17.4 88.9 9.8 14.3 0.2 27.3 0.2 Morella cerifera (L.) Small Wax Myrtle 14.3 0.1 Subtotal 25.4 35.2 57.4 62.6 Leaves and stems Morella cerifera Wax Myrtle 83.3 7.4 88.9 16.1 28.6 0.9 72.7 7.1 Phoradendron serotinum Mistletoe 33.3 2.1 77.8 7.1 18.2 0.1 Quercus sp. Oak 55.6 2.8 83.3 3.7 42.9 0.1 45.5 0.3 Ilex vomitoria Yaupon Holly 22.2 1.4 61.1 5.4 14.3 0.6 36.4 0.2 Vitis sp. Grape 72.2 6.9 22.2 1.3 Salicornia sp. Glasswort 66.7 3.3 42.9 3.0 27.3 0.3 Zanthoxylum clava-herculis L. Hercules Club 22.2 2.6 55.6 3.0 42.9 0.9 Juniperus virginiana L. Redcedar 33.3 2.1 16.7 0.9 14.3 0.2 18.2 0.1 Prunus caroliniana (P. Mill.) Ait. Cherrylaurel 22.2 3.1 33.3 0.4 14.3 0.3 Pinus sp. Pine 5.6 0.1 11.1 0.1 18.2 0.1 Smilax sp. Greenbriar 16.7 0.8 50.0 1.6 14.3 0.3 9.1 0.1 Persea borbonia (L.) Spreng. Redbay 18.2 0.1 2009 B.W. Morse, M.L. McElroy, and K.V. Miller 583 Oct–Dec 2004 Jan–Feb 2005 Oct–Dec 2005 Jan–Mar 2006 n = 18 n = 18 n = 7 n = 11 Food item (species) Common name % Occ. Vol. % Occ. Vol. % Occ. Vol. % Occ. Vol. Leaves and stems Salix sp. Willow 18.2 0.1 Baccharis halimifolia L. Groundsel Bush 9.1 0.1 Vaccinium arboreum Marsh. Sparkleberry 14.3 0.1 18.2 0.1 Unknown stem 71.4 8.8 54.5 11.3 Unknown leaves 42.9 0.9 45.5 0.4 Subtotal 29.3 42.9 16.1 20.4 Forbs Polygonum sp. Smartweed 27.8 2.9 38.9 1.0 45.5 0.2 Pleopeltis polypodioides (L.) Andrews & Windham Resurrection Fern 16.7 0.2 66.7 5.5 14.3 0.1 90.9 6.8 Tillandsia usneoides L. Spanish Moss 33.3 9.6 55.6 2.1 100.0 8.1 Unknown I 14.3 0.1 18.2 0.3 Unknown II 9.1 0.1 Subtotal 12.6 8.6 8.3 7.4 Grass and Grasslike 88.9 26.1 55.6 2.0 100.0 14.9 81.8 8.1 Fungi/Lichen 16.7 0.4 66.7 7.3 42.9 0.8 54.5 0.5 Other 5.6 0.1 14.3 0.2 45.5 0.6 Unidentified 100.0 6.2 100.0 4.0 100.0 2.6 100.0 0.8 584 Southeastern Naturalist Vol. 8, No. 4 Appendix 2. Percent relative density of food items in feces of Fallow Deer on Little St. Simons Island, GA, 2004–2005 as determined by microcompositional analysis. Fall 2004 Winter 2005 Spring 2005 Summer 2005 Fall 2005 Plant (species) Common name n = 12 n = 14 n = 13 n = 12 n = 17 Grass and Grasslike Andropogon sp. Broomsedge 0.4 Andropogon glomeratus (Walt.) B.S.P. Bushy Broomsedge 0.3 Andropogon virginicus L. Broomsedge Bluestem 0.3 0.7 0.3 Aristida sp. Three-awn Grass 0.4 Cynodon dactylon (L.) Pers. Bermudagrass 5.4 0.6 4.7 4.3 7.8 Cyperus spp. Sedge 1.4 2.8 5.3 8.3 6.4 Distichlis spicata (L.) Greene Saltgrass 6.8 9.5 8.8 19.2 4.7 Eleocharis sp. Spike-rush 0.9 0.4 Eragrostis sp. Lovegrass 9.0 1.5 4.4 1.4 0.8 Festuca sp. Fescue 0.4 Fimbristylis sp. Fimbry 1.6 0.7 Juncus sp. Rush 0.7 4.0 0.9 0.7 0.8 Piptochaetium avenaceum (L.) Parodi Blackseed Speargrass 0.6 0.3 3.3 Rhynchospora sp. Beaked-rush 0.7 0.3 Setaria sp. Foxtail 0.3 0.7 Setaria viridis (L.) Beauv. Green Bristlegrass 0.4 0.6 0.7 0.8 Spartina sp. Cordgrass 0.4 0.9 1.9 1.8 Sporobolus sp. Dropseed 0.3 1.3 0.4 0.6 Sporobolus virginicus (L.) Kunth Coastal Dropseed 0.3 0.4 Stenotaphrum sp. St. Augustine Grass 0.3 Triplasis purpurea (Walt.) Chapman Purple Sandgrass 0.3 0.7 Unolia sp. Sea Oats/Wood Oats 30.6 9.5 23.9 19.9 13.1 Unknown grass I 0.3 1.1 0.3 Unknown grass II 0.7 2.2 0.6 1.1 0.6 Subtotal 56.1 32.6 56.9 67.0 36.6 2009 B.W. Morse, M.L. McElroy, and K.V. Miller 585 Fall 2004 Winter 2005 Spring 2005 Summer 2005 Fall 2005 Plant (species) Common name n = 12 n = 14 n = 13 n = 12 n = 17 Browse Acer sp. Maple 1.4 1.2 1.9 Bumelia sp. Buckthorn 0.4 0.6 0.7 1.7 Callicarpa americana L. American Beautyberry 0.3 1.1 Celtis sp. Hackberry 1.4 0.4 Hibiscus sp. Mallow 1.1 Juniperus virginiana L. Redcedar 1.4 5.5 3.8 1.1 0.8 Kosteletzkya virginica (L.) K. Presl ex Gray Seashore Mallow 0.4 1.1 Magnolia grandifl ora L. Southern Magnolia 0.6 Osmanthus sp. Wild-olive 0.9 0.7 0.6 Persea sp. Bay 0.7 1.2 2.2 Persea borbonia (L.) Spreng. Redbay 3.1 0.6 Pinus sp. Pine 0.6 2.8 0.4 0.8 Pinus taeda L. Loblolly Pine 0.3 Quercus spp. Oak 19.8 27.1 8.5 2.5 13.4 Rhododendron type 0.3 Rhus sp. Sumac 0.3 Rubus sp. Blackberry 4.0 0.0 0.3 Salix sp. Willow 2.2 4.9 1.3 1.4 3.4 Vaccinium sp. Blueberry 0.7 1.6 Zanthoxylum clava-herculis L. Hercules' Club 0.8 Subtotal 28.1 49.8 23.9 9.8 22.9 Forbs Bidens sp. Spanish Needles 0.3 Chenopodium sp. Mexican Tea 0.4 0.3 0.4 0.3 Croton punctatus Jacq. Gulf Croton 1.8 5.5 1.3 1.4 1.4 Ipomoea sp. Morning Glory 0.7 0.8 Lepidium virginicum L. Peppergrass 0.9 1.1 Limonium carolinianum (Walt.) Britt. Sea-lavender 0.4 586 Southeastern Naturalist Vol. 8, No. 4 Fall 2004 Winter 2005 Spring 2005 Summer 2005 Fall 2005 Plant (species) Common name n = 12 n = 14 n = 13 n = 12 n = 17 Forbs Oenothera sp. Evening Primrose 0.3 Oxalis sp. Woodsorrel 0.6 Polygonum sp. Smartweed 0.6 0.4 Salicornia sp. Glasswort 0.3 0.3 Salivia sp. Sage 0.3 Salsola sp. Thistle 0.7 Salsola kali Russian Thistle 0.3 Solidago sp. Goldenrod 0.4 Suaeda linearis (Ell.) Moq. Sea-blite 0.3 Tradescantia sp. Spiderwort 0.4 0.9 Typha latifolia Cattail 0.4 0.6 0.3 Verbascum sp. Mullein 0.3 0.3 Unknown forb I 3.6 4.6 1.3 1.1 20.1 Unknown forb II 6.3 6.9 0.3 Unknown forb III 1.4 Composite 0.6 0.3 Subtotal 7.2 12.6 12.6 13.0 25.7 Other Zea mays L. (Corn) kernel 1.6 Flower 0.6 Grass seed and glume 4.0 4.6 2.8 6.9 3.6 Legume pod 0.4 1.1 0.3 Seed 4.3 0.6 2.2 1.8 10.3 Cetraria sp. Fungi 0.4 Subtotal 8.6 5.2 6.6 10.1 14.8