2012 SOUTHEASTERN NATURALIST 11(3):487–506 Food of the Armadillo Dasypus novemcinctus L. from Cumberland Island, GA John O. Whitaker, Jr.,1,* Carol Ruckdeschel2, and Laura Bakken1 Abstract - We provide a review of Dasypus novemcinctus (Nine-banded Armadillo) food studies and report on the diet on Cumberland Island, GA. Major invertebrate foods eaten by Armadillos on Cumberland Island were ants (19.3%); beetles, adults and larvae (27.4 %); centipedes (6.5%); lepidopterans, mostly larvae (caterpillars, 13.2%); millipedes (5.7%); spiders (2.1%); and sowbugs (2.2%). Vertebrates eaten (1.9% volume) were mostly frogs and lizards. Plant material comprised about 12% of the volume, including much fruit such as Melia azedarach (China Berry), Vitis (grapes), Ampelopsis arborea (Pepper Vine), and Serenoa repens (Saw Palmetto). The subterranean fungi, Endogonaceae, were eaten by 23.9% of the Armadillos, comprising 1.6% of the volume of their diet. The millipede Narceus sp. appears to have been greatly reduced, based on the significant reduction of this species as a component of the gut content over three decades. This reduction could be due to Armadillo predation. Ants, scarabaeid adults and larvae, and spiders also showed significant decreases during the period of study, which could be a result of Armadillo predation. A number of items—elaterid larvae, crickets, caterpillars, centipedes, sowbugs, and Endogonaceae—showed significant increases as components of Armadillo stomachs, but it is not known whether these changes might be related to Armadillo predation. The increases could have been due to other causes such as habitat or climatic changes. Introduction There is relatively little detailed information on the food of Dasypus novemcinctus L. (Nine-banded Armadillo; hereafter Armadillo) from barrier islands or maritime forest ecosystems. Carol Ruckdeschel has been studying the ecology of Cumberland Island since the early 1970s, and this paper constitutes a part of that study. Armadillos are considered exotic in Georgia, and were unrecorded in the US before 1849 (Humphrey 1974). They arrived in Georgia from Mexico and south Texas, and from a few animals which escaped confinement in Florida. The Armadillo roots in the ground litter, pig fashion, by pushing forward with its nose along the ground. We suspect that much of its food is detected by olfaction. How much it uses its eyes and takes items off the surface is unknown. Earthworms, ants, caterpillars, beetle larvae, Endogonaceae, and probably many other foods are likely found by odor. Although we do not have estimates of availability of the soil organisms, we suspect that the diets of Armadillos may give a reasonable estimate of available soil organisms. 1Department of Biology, Indiana State University, Terre Haute, IN 47809. 2Cumberland Island Museum, Cumberland Island, PO Box 7, St. Marys, GA 31558. *Corresponding author - John.Whitaker@indstate.edu. 488 Southeastern Naturalist Vol. 11, No. 3 The Armadillo arrived on Cumberland Island, Camden County, in approximately 1973. The species was successful, increasing in numbers rapidly, and is now abundant. It is a major predator on the invertebrates of the island, while it has few predators, perhaps mainly Bubo virginiana Gmelin (Great Horned Owl) and Alligator mississippiensis (Daudin) (American Alligator), and to a lesser extent Lynx rufus (Schreber) (Bobcat) and Canis latrans Say (Coyote). One can usually see several Armadillos as one walks along any of the trails in early morning, late afternoon, or at night in summer, or at midday in winter. Osborn et al. (2000) examined stomachs of 171 Armadillos from Cumberland Island but identified only 18 foods. Major foods in all seasons were beetle adults (especially Elateridae, Scarabaeidae, and Cicindelidae) and larvae (especially Scarabaeidae and Elateridae), ants and wasps (all stages), and lepidopteran larvae and pupae. Fruit was important only in August, the most important being Vitus (grapes) and Serenoa repens (W. Bartram) Small (Saw Palmetto), but Smilax sp. (greenbriar) and Prunus caroliniana (P. Mill.) Ait. (Carolina Laurelcherry) were also eaten. Vertebrate prey included amphibians (Scaphiopus holbrookii (Harlan) [Eastern Spadefoot]) and reptiles (Eumeces sp., Anolis carolinensis (Voigt) [Carolina Anole], Sceloporus undulatus (Bosc and Daudin in Sonnini and Latreille) [Eastern Fence Lizard], Opheodrys aestivus (L.) [Rough Greensnake]) and their eggs. Kalmbach (1943) recorded 488 different food items from 169 stomachs of Armadillos from Texas, mostly from March through July. Insects formed 77.6% of the food items and 41.6% were beetles, the majority being adult and larval scarabaeids. Carabids accounted for 9.4% of the total volume, and were eaten by all but a few individuals. Hymenopterans, mostly ants, were the second most important order, forming 14.0% of the diet, and nearly every stomach contained ants. One individual had eaten 40,000 ants of 3 species. Wasps, velvet “ants” (Mutillidae), digger wasps (Scoliidae), and paper wasps (Vespidae) occurred occasionally. Caterpillars (mostly Noctuidae and Notodontidae) occurred in 145 (85.7%) of the 169 stomachs, moths in only 10. Orthopterans (6.2%), most commonly roaches (Blattellidae, Ischnoptera sp.), but also including crickets, grasshoppers, and walking sticks, were eaten by all but 12 individuals. Termites (4.5%) were consumed in large numbers by some individuals, and occurred in 126 (74.5%) stomachs. Hemiptera (2.0%) were eaten by 128 (75.7%) of the Armadillos. Dipteran larvae accounted for 1.5% of the volume. Arachnids included spiders, scorpions, solpugids, ticks, and mites. Millipedes and centipedes together accounted for 6.2% of the volume, and were eaten by 89.3% of the 169 Armadillos examined. Earthworms, most important in January, formed most of the 6.2% volume of “miscellaneous invertebrates”. Crayfish, isopods, slugs, and snails were occasionally eaten. Reptiles and amphibians formed 1.2% of the volume, but few Armadillos were taken in winter, when Wirtz et al. (1985) and Sikes et al. (1990) found them regularly eaten. Few birds or their eggs were eaten. Rubus (blackberries), 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 489 Morus (mulberries), Callicarpa (French mulberries), Prunus (plums), and Ilex (holly) were eaten, and mushrooms and puffballs were heavily eaten by a few individuals. Smith et al. (1998) found chitinase activity in this species, which may aid in digestion of fungi, as well as insects and other arthropods. Wirtz et al. (1985) examined 186 Armadillo stomachs from Florida and identifi ed over 167,000 individual food items, more than half being ants. Insect larvae were the dominant food throughout the year. Beetles were heavily eaten, with 25 families being found. The larvae of Elateridae and Scarabaeidae were most important. Lepidopteran larvae, Orthoptera, and Diptera were important in all seasons, and Homoptera, Hemiptera, Isoptera, and Dermaptera were eaten occasionally. Spiders, centipedes, millipedes, and earthworms were also consumed. Lizards and anurans were often eaten, especially in winter, when they were presumably torpid and thus easy prey. Nesbitt et al. (1979) examined stomachs from 172 Armadillos from Florida and found insects comprising 78.5% by volume, with beetles, especially scarabaeids, the largest component (29.7%). Hymenoptera (15.1%), Diptera (13.3%), and Orthoptera (10.5%) were also important. Some economically important insects found in these stomachs were Spodoptera frugiperda (J.E. Smith) (Fall Armyworm), Pycnocelus surinamensis L. (Surinam Cockroach), and eggs of Romalea microptera (Beauvois) (Lubber Grasshopper). Layne (1976) was concerned that Armadillos might be a threat to Florida’s rare endemic reptiles, noting that Sceloporus woodi Stejneger (Florida Scrub Lizard), Neoseps reynoldsi Stejneger (Sand Skink), Rhineura floridana (Baird) (Florida Worm Lizard), and Tantilla relicta Telford (Florida Crowned Snake) have been found in Armadillo stomachs at the Archbold Biological Station. Staller et al. (2005) found that the Armadillo was a major predator on eggs of Colinus virginianus L. (Northern Bobwhite) in Florida and Georgia, and Douglass and Winegarner (1977) found evidence that it commonly feeds on eggs of Gopherus polyphemus (Daudin) (Gopher Tortoise) in southern Florida. Fitch et al. (1952) found that the main foods of the Armadillo in Louisiana were beetles, but important secondary foods were orthopterans and reptiles in winter, lepidopteran and dipteran larvae in spring, and fruits in autumn. Newman and Baker (1942) reported an Armadillo eating three young Sylvilagus floridanus (J.A. Allen) (Eastern Cottontail). Hamilton (1946) found the food of eight Armadillos in summer to be 80% Diospyros virginiana L. (Black Persimmon) fruit. Taber (1945) found that Armadillos readily fed on maggotinfested chickens but were slow to feed on fresh chickens. Clark (1951) observed Armadillos tearing up paper nests of wasps, apparently to obtain the larvae and pupae. The purpose of the present study was to determine the diet of the Armadillo over time on Cumberland Island, and to compare the results with other studies. 490 Southeastern Naturalist Vol. 11, No. 3 Materials and Methods The Armadillos were collected by Carol Ruckdeschel and C.R. Shoop. A total of 134 Armadillo stomachs was available, including samples from every month: January (12), February (6), March (6), April (15), May (26), June (23), July (6), August (10), September (15), October (6), November (7), and December (2), in the years from 1975 through 2006. Most of the Armadillos were found dead on the road or dead on the beach, and others were taken as nuisance animals. The stomach was immediately removed from each Armadillo and preserved in 70% alcohol, then sent to J.O. Whitaker, Jr. for analysis. Original examination of the contents of a stomach generally took about one day each. The stomach contents were put into a 0.8-mm mesh strainer to remove detritus and small particles. The strained material was occasionally examined to be sure that whole small organisms were not passing through the strainer. The items were then sorted and counted, with different species or species groups (such as ants) being put into different vials for further identification to smaller taxonomic groups. The percent volume of each item in each stomach was estimated visually. It was noted that many of the items were whole or nearly so, even though they had passed through the mouth and had presumably been processed by the teeth before entering the stomachs. The data were summarized by adding all of the individual values for each item. Those sums were divided by the total number of stomachs x 100 for each month and for the overall estimate of percent volume during the study. Percent frequency was calculated for each item as the percentage of the total number of stomachs in which a given item occurred. The amounts of the various food items were presented as total numbers and percentages. Description of Study Area Cumberland Island is the largest and southernmost of the barrier islands off the coast of Georgia. It was designated a National Seashore in 1972, and part of the island was established as a Wilderness Area in 1982. The island is about 27 km (17 miles) long and averages about 2.4 km wide. A few residents and one active hotel were present before the Wilderness designation, and the National Park Service maintains a few buildings and campgrounds. There is one road running the length of the island, with several east–west roads connecting habitats, from grassy temporary wetlands and dunes to upland maritime forest. The island is surrounded by saltwater, but there is fresh groundwater from rainwater retention. There are some small ponds, creeks, swamps, and marshes, and the largest natural freshwater lake occurring on any of the Georgia barrier islands. The major portion of the island is maritime forest, with various live oaks, hickories, and pines. Tillandsia usneoides (L.) L. (Spanish Moss) is abundant on the hardwoods and would litter the ground if it were not for the feral livestock. Sabal palmetto (Walt.) Lodd. (Cabbage Palm) and Saw Palmetto are very common in the understory of the upland. 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 491 Feral horses and swine on Cumberland Island run freely, thus tramping down the soil and litter and having a great impact on island ecology. Several mammal species that are likely limited by the feeding and tramping of these animals are shrews, arvicoline rodents, and Sigmodon hispidus Say and Ord (Cotton Rat). Similar tramping has been shown to be detrimental to small mammals in Indiana (Whitaker 1967). The droppings from feral livestock do probably provide fertilizer and nurture a food chain that would benefit some species. The heavy tramping and rooting probably limits the available food for Armadillos as well. Results and Discussion Major food items eaten by Armadillos from Cumberland Island (Appendices 1, 2) were beetles (adults and larvae; 27.4% of total food volume), ants (19.3%), caterpillars (11.5%), centipedes (6.5%), millipedes (5.6%), spiders (2.1%), and sowbugs (2.2%). A few vertebrates were eaten (1.9% volume), mostly lizards and their eggs. Various plant materials were occasionally eaten, principally fruit. Fungi in the Family Endogonaceae were often eaten, totaling 1.6% of the volume. Many of the foods, even larger ones, were not chewed much, such as centipedes (6 or 7 cm in length), large lepidopteran larvae and pupae, and large beetle larvae. In contrast, nearly all of the hundreds of adult beetles were chewed and very few left whole. Large millipedes, Narceus sp., were cut into chunks. Insects Beetles and beetle larvae were taken throughout the year and included many families (14 identified) and species. Particularly obvious was the large carabid beetle, Calosoma scrutator Fabr. (Fiery Searcher). It appeared in stomachs in every month, April through August, but was particularly abundant in May. Adults of this species formed 2.4% of the volume of food over the study. Its larvae were found in May, June, and July, and they formed 2.3% of the total volume. Other carabids (412 individuals) were found in every month, and made up 2.9% of the total volume. Scarabaeid beetles were found in stomachs in all months except December (n = 2), and were especially abundant April through June and again August through October. A total of 1547 adult individuals were found, and they were included in 55.2% of the stomachs. Their larvae (grub worms) were also abundant, especially during May through October. Adult click beetles (Elateridae) were found in every month, and their larvae (wireworms) were heavily eaten in all months. A few long-horned beetles (Cerambycidae) were found, mostly in August. Numerous unidentified coleopteran larvae occurred, especially from January through April, forming 0.9% of the total volume. A moderate number of rove beetles (Staphylinidae) were in stomachs, the greatest number in May. Tenebrionids were commonly found, especially in August through October. 492 Southeastern Naturalist Vol. 11, No. 3 Ants (Formicidae, Hymenoptera) were the most abundant organisms found, with over 23,000 found in the 134 stomachs (x̅ = about 175 per stomach). They formed 19.3% of the volume of food, and occurred in 128 of the stomachs (95.5%). These included a large number of species. The ants were often obtained from nests, since all stages of ants, including eggs, were represented in stomach contents. A few stinging hymenopterans, including vespid wasps and bees, were found. Caterpillars (lepidopteran larvae) were heavily eaten by Armadillos in every month of the year (x̅ = about 10 per stomach), formed about 11.5% of the volume, and were found in 114 (85%) of the stomachs. Lepidopteran pupae occurred in 22% of the stomachs and formed 1.0% of the volume. A total of 2539 larvae, 113 pupae, and 338 adult moths were found. Termites (Isoptera) were also eaten, especially from January through April. These were obviously obtained from termite nests, as indicated by the large numbers and various stages present. Orthopterans eaten were mostly crickets, including many field crickets (Gryllus spp.) and a few little brown crickets (Nemobius spp.) Gryllus formed 3.1% of the volume, occurring in 43.3% of the stomachs, and included 157 individuals. Hemipterans and homopterans were found, but none were very abundant. Relatively few dipterans were found, the most abundant being midge adults and larvae (Chironomidae), cranefly larvae (Tipulidae), and unidentified larvae. The Armadillos fed little on grasshoppers or dragonflies, presumably because they are difficult to capture. Other invertebrates Centipedes (Chilopoda) and millipedes (Diplopoda) were heavily eaten throughout the year. At least 3 species of centipedes were included and collectively formed 6.5% of the food, were found in 78.4% of the Armadillos, and were represented by 1019 individuals (x̅ = 7.6). There were also three species of millipedes: a very large, dark-colored Narceus sp.; a small, dark coiled one; and a flattened, colorful one. Many of the items were eaten mostly intact, including all of the centipedes and millipedes except for Narceus sp., which was cut into chunks before swallowing. Millipedes (including Narceus sp.) comprised 5.6% of the volume, and were represented by 1296 individuals (x̅ = 9.6). Narceus sp. was found in 17.2% of the stomachs, other millipedes in 62.7% of stomachs. Spiders occurred in relatively small numbers, but were present throughout the year, forming 2.1% of the volume, with a total of 226 spiders in 59% of the the stomachs. Sowbugs (3 species) occurred in every month but mostly in January through June. Earthworms and snails occurred in relatively small numbers. Earthworms were found in 15.7% of the stomachs, comprised 0.6% of the volume, and included 220 individuals. Vertebrates Relatively few vertebrates were eaten by Armadillos on Cumberland Island, forming less than 2% of the volume. Various lizards and lizard eggs were the 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 493 most common, including anoles and skinks, followed by frogs and toads. Two clutches of lizard eggs were found—probably of Sceloporus undulatus (Bosc & Daudin) (Eastern Fence Lizard)—and most of the eggs were intact. Bird remains were found in a few stomachs, and mammal remains in one. The vertebrates, of course, may have been taken as carrion. Plant materials and fungi A variety of plant material was found, including various roots, leaves, fruit, and seeds. Some of the fruits identified were Melia azedarach L. (China Berry, 0.4%); Vitus (grapes) and Ampelopsis arborea (L.) Koehne. (Pepper Vine) (0.1%); and Diospyros virginiana L. (Persimmon, 6%). Fungal foods included mushrooms and the tiny subterranean fungi in the Family Endogonaceae. Even though the clumps (n = 133) are tiny, they comprised 1.6% of volume, and were found in 32 of the stomachs. Seasonal differences Various beetles, ants, and caterpillars were found throughout the year (Appendix 1). Larger numbers were often taken in summer, such as with ants, of which the largest numbers were eaten in April and summer. Centipedes, millipedes, sowbugs, and spiders also were eaten almost throughout the year. The lists of foods eaten by month are given in Appendix 1. The carabid larvae and adults were taken from April through July. Some amphibians and reptiles were taken in winter, with 6 being taken in January, 5 in February, and most of the rest in April and May. Lizard eggs were eaten from April through September. How do foods eaten by Armadillos on Cumberland Island compare to foods eaten elsewhere by this species? The total volume of insects eaten by Armadillos was 68.4% on Cumberland Island (this study), 77.6% in Texas (Kalmbach 1943), and 78.5% in Florida (Nesbitt et al. 1979). Beetles and their larvae and ants were the insects most often eaten: beetles 27.4% and ants 19.3% on Cumberland Island, 41.6% and 14.3%, respectively, in Texas. The most important beetles in most studies were scarabaeids, elaterids, and carabids. Ants were found in almost every stomach from Cumberland Island and in Florida (Wirtz et al. 1985) and comprised over half the individuals in both the areas. Caterpillars were important diet items on Cumberland and from Florida, and were found in 85.7% of the stomachs from Texas, but adult moths made up little of the food in any of the studies. Orthopterans, mostly crickets and roaches, made up 5.1% of the volume of food on Cumberland Island, 6.2% of the food volume in Texas, and were major items in Florida (Wirtz et al. 1985). Millipedes and centipedes made up 9.1 and 2.7%, respectively, of the food on Cumberland Island, totaled 6.2% in Texas (Kalmbach 1943), and were important in Florida (Wirtz et al. 1985). Reptiles and amphibians, mostly lizards, formed a small amount of the food in nearly every study. Plant materials and fungi comprised 12% of the volume of food on Cumberland Island and made up small amounts in the rest of the study areas. 494 Southeastern Naturalist Vol. 11, No. 3 Armadillos feed on a variety of available items, and there is great similarity in the items of foods heavily eaten in the various study areas. There was no major item eaten on Cumberland Island that was not eaten elsewhere. Fungi of the family Endogonaceae formed 1.6% of the diet on Cumberland Island, but were not reported elsewhere. However, fungi are overlooked by most workers. The large carabid beetle, Calosoma scrutator, was apparently sharply declining from the 1980s through the turn of the century (this study) and was not found by Osborn et al. (2000) on Cumberland, further indicating it has nearly disappeared from Cumberland. Also it has not been previously reported from Camden County. There is much similarity in the foods from various regions and relatively few major differences. Impact of Armadillos on the invertebrate community To determine whether predation by Armadillos has resulted in changes in abundances of prey species, we divided the food data into decades, primarily the 1980s, 1990s, and 2000s. Three stomachs from the 1960s and 1970s were included with data from the 1980s. We examined the items in Appendix 2 and selected those that were large enough in sample size to test. Using the data in Appendix 2, we calculated estimated volumes that would be expected based on the percent of stomachs from each decade (Table 1). If there was no change, then the actual volumes should be similar to the expected numbers. Major differences would suggest changes in invertebrate populations between the decades. Differences were tested for signifi- cance using chi-square. Such differences, of course, might be due to factors other than Armadillo predation, such as climatological events, vegetative community Table 1. Expected and actual values for numbers of individuals of selected foods of Nine-banded Armadillo on Cumberland Island, GA. 1980s 1990s 2000s Total No. stomachs examined 57 31 46 134 % of stomachs 42.5 23.1 34.3 Prey Expected Actual Expected Actual Expected Actual Scarab adult 247 307 134 153 199 121 Scarab larvae 187 252 102 50 151 139 Calosoma scrutator adult 132 299 72 24 107 0 Calosoma scrutator larvae 131 263 24 48 106 1 Elateridae larvae 290 188 159 165 236 336 Ants 11,053 12,800 6011 9500 8920 3684 Gryllid larvae 180 121 98 97 145 205 Lepidoptera 654 538 356 422 528 579 Chilopoda 369 377 201 127 298 364 Narceus sp. 103 168 56 70 83 5 Diplopoda 216 104 118 85 175 320 Sowbug 105 95 57 26 84 125 Earthworm 61 65 33 7 49 71 Endogonaceae 89 40 49 73 72 97 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 495 changes such as those resulting from the listing as a National Seashore, and cumulative changes resulting from feral livestock, or to the presence of the fire ant. The most obvious change between decades was that of the millipede Narceus sp. Its numbers observed over the three decades were 168, 70, and 5 (Table 1). Expected values based on the numbers examined in the three decades were 103, 56, and 83, and these differences were significant (chi-square = 117.8). It is possible that the reduced numbers through time were due to the Armadillos, as these millipedes are large and often conspicuous. Also interesting was that the other millipedes were taken in significantly greater numbers (chi-square = 187.37) over the three decades, 104, 85, and 320 (expected values 216, 118, and 175). We do not know the ecological relationships between Narceus sp. and the other millipedes, but it is possible that the decrease in Narceus sp. led to the increase in the other millipedes. Of the other forms tested, ants (31.45), scarabaeid adults (47.83), scarabaeid larvae (50.05), and spiders (23.8) showed significant decreases in numbers (respective chi-square values in parentheses), which may be a result of Armadillo predation. Ants are abundant, and since ants are the major food of Armadillos on Cumberland Island, it is logical that they might have diminished, although it would be difficult to prove that the decrease was because of Armadillos. There are relatively few earthworms on Cumberland Island, with only 65, 7, and 51 being taken over the three decades as compared to expected numbers of 52, 28, and 42. Relatively greater numbers of earthworms were eaten during the middle rather than in the earlier or later decades. These numbers were signifi cantly different from random (chi-square = 85.6). Perhaps earthworms were increasing on Cumberland Island until the advent of the Armadillos, and the Armadillos might at that time have begun to have a negative influence on them, but this does not explain their increase in the third decade. Perhaps climatologic conditions could have played a role. Other items—wireworms (elaterid larvae) (81.22), crickets (44.14), caterpillars (27.1), centipedes (42.03), sowbugs (37.8), and the underground fungi Endogonaceae (117.8)—all showed significant increases. It is unclear how increases in any of these invertebrates might have been related to Armadillos. Contrary to what happened in other groups, the larval elaterids increased, whereas the adults showed no change. This is hard to explain. However, there is a large number of species in this family, and adults are not eaten in large numbers (only 118 in stomachs in the three decades). Perhaps many of the adults are in different habitats and thus less available to the Armadillos, or it may be, as indicated by the small numbers eaten, that adult click beetles are not as appealing to the Armadillos as food as are the larvae. Fungi of the family Endogonaceae look like tiny clods of dirt at low power of the dissecting scope, and are probably often overlooked. However, at powers above 40x they look like tiny clusters of grapes, with each sporocarp being stalked. It was surprising to find these fungi being eaten by Armadillos, as they are so small. It might be suspected that they were taken incidentally 496 Southeastern Naturalist Vol. 11, No. 3 with some other food item. However, they are heavily eaten and presumably desired by some small mammals. For example, Endogonaceae formed about 16.5% of the diet of Zapus hudsonius Zimmermann (Meadow Jumping Mouse) and about 33.3% of the diet of Napaeozapus insignis (Miller) (Woodland Jumping Mouse), both in central New York (Whitaker 1963a, b). Also it formed about 3.8% of the diet of Blarina carolinensis (Bachman) (Southern Short-tailed Shrew) on Cumberland Island (Whitaker and Ruckdeschel 2006). These species clearly seek it as food, as sometimes as much as 100% of their food is of this item. We suspect that the Armadillo found this food by scent since we see no other food with which it might be ingested, and since it was taken so often. These fungi produce odor attractive to mammals, which then disperse the spores through defecation. Therefore it is logical to think that these fungi may have increased by being eaten and their spores dispersed by Armadillos, especially as the Armadillo is more wide-ranging than the Southern Short-tailed Shrew. Many kinds of ground beetles, Carabidae, were eaten, and populations of these showed no significant difference over the three decades (chi-square = 1.78), except for Calosoma scrutator. It is large (25–36 mm) and showy, having the head, prothorax, and elytra blackish but edged with greenish and gold. Both adults and larvae decreased radically during this study. In the first decade, 299 adults were eaten, 24 in the second decade, and none in the third decade, as opposed to 132, 72, and 107 expected. This difference was significant (chi-square = 12,548). Correspondingly, 263, 48, and 1 larvae of this species were found among the Armadillo prey in the three decades, as opposed to expected numbers of 131, 24, and 106. This difference also was significant (chi-square = 243). We suspect that numbers of this interesting carabid could have been reduced by the Armadillo over this period. We conclude that Armadillos are relatively unselective feeders and that information on their food gives a fairly good sampling of the soil invertebrates on Cumberland Island. This, in turn, can suggest whether foods are increasing, decreasing, or remaining stable. In addition, population sizes of some of their prey may be influenced by Armadillos, although there may be other factors influencing them, such as (1) feral livestock—pigs and horses; (2) the current long-term drought in southern Georgia; and (3) the recent arrival of fire ants, whose relation to insects and other biota is unknown. In some areas, Armadillos can cause damage such as destruction of quail eggs and poultry, burrowing damage to structures, damage to crops and gardens, and predation on reptiles and amphibians. On Cumberland Island, many of the perceived negative effects are unlikely because most of the area is in Wilderness. However, coupled with feral swine and horses, they could impact shorebird nesting. Possible good effects of Armadillos are that their burrows can be used by other animals such as Alligator mississippiensis (Daudin) (American Alligator), rattlesnakes, opossums, cottontails, skunks, Neovison vison (Schreber) 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 497 (American Mink), and Ondatra zibethicus (L.) (Muskrat), and that they stir and aerate the soil. Acknowledgments Linda Castor did most of the calculations. C. Robert Shoop helped with the collection of Armadillos. Literature Cited Clark, W.K. 1951. Ecological life history of the armadillo in the eastern Edwards Plateau region. American Midland Naturalist 46:337–358. Douglass, J.F., and C.E. Winegarner. 1977. Predators of eggs and young of the Gopher Tortoise, Gopherus polyphemus (Reptilia, Testudines, Testudinidae) in southern Florida. Journal of Herpetology 11(2):236–238. Fitch, H.S., P. Goodrum, and C. Newman. 1952. The armadillo in the southeastern United States. Journal of Mammalogy 33:21–37. Hamilton, W.J., Jr. 1946. The Black Persimmon as a summer food of the Texas armadillo. Journal of Mammalogy 27:175. Humphrey, S.R. 1974. Zoogeography of the Nine-banded Armadillo (Dasypus novencinctus) in the United States. Bioscience 24(8):457–462. Kalmbach, E.R. 1943. The armadillo: Its relation to agriculture and game. Game, Fish, and Oyster Commission, Austin, TX. 61 pp. Layne, J.N. 1976. The armadillo, one of Florida’s oddest animals. Florida Naturalist 49:8–12. Nesbitt, S.A., W.M. Hetrick, L.E. Williams, Jr., and D.H. Austin. 1979. Foods of the Nine-banded Armadillo in Florida. Proceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies 31:57–61. Newman, C.C., and R.H. Baker. 1942. Armadillo eats young rabbits. Journal of Mammalogy 23:450. Osborn, D.A., M.I. Nelson, and R.J. Warren. 2000. Armadillo diets among seasons and between habitats on Cumberland Island, Georgia. Proceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies 54:282–291. Sikes, R.S., G.A. Heidt, and D.A. Elrod. 1990. Seasonal diets of the Nine-banded Armadillo (Dasypus novemcinctus) in a northern part of its range. American Midland Naturalist 123:383–389. Smith, S.A., L.W. Robbins, and J.G. Steiert. 1998. Isolation and characterization of a chitinase from the Nine-banded Armadillo, Dasypus novemcinctus. Journal of Mammalogy 79:486–491. Staller, E L., W.E. Palmer, J.P. Carroll, R.P. Thornton, and D.C. Sisson. 2005. Identifying predators at Northern Bobwhite nests. Journal of Wildlife Management 69:124–132. Taber, F.V. 1945. Contributions on the life history and ecology of the Nine-banded Armadillo. Journal of Mammalogy 26:211–226. Whitaker, J.O., Jr. 1963a. A study of the Meadow Jumping Mouse, Zapus hudsonius Zimmermann, in central New York. Ecological Monographs 33:215–154. Whitaker, J.O., Jr. 1963b. Food, habitat and parasites of the Woodland Jumping Mouse in central New York. Journal of Mammalogy 44:316–321. 498 Southeastern Naturalist Vol. 11, No. 3 Whitaker, J.O., Jr. 1967. Habitat and reproduction of some of the small mammals of Vigo County, Indiana, with a list of the mammals known to occur there. Occasional Papers of the C.C. Adams Center for Ecological Studies. No. 16. 24 pp. Whitaker, J.O., Jr., and C. Ruckdeschel. 2006. Food of the Southern Short-tailed Shrew (Blarina carolinensis) on Cumberland Island, Georgia. Southeastern Naturalist 5:361–366. Wirtz, W.O., D.H. Austin, and G.W. Dekle. 1985. Food habits of the common Long-nosed Armadillo (Dasypus novemcinctus) in Florida. Pp. 439–451, In G.G. Montgomery (Ed.). The Evolution and Ecology of Armadillos, Sloths, and Vermilinguas. Smithsonian Institution Press, Washington, DC. 451 pp. 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 499 Appendix 1. Foods (% volume, frequency) eaten by 134 Nine-banded Armadillos on Cumberland Island, GA, with summarized data for all stomachs (total). Percent volume Frequency Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Total Percent n 12 6 6 15 26 23 6 10 15 6 7 2 134 Coleoptera (27.4) Scarabaeidae – adult 1.8 2.3 0.5 6.4 7.4 4.5 0.5 3.8 4.9 5.3 0.7 0 4.3 74 55.2 Scarabaeidae – larvae 2.2 0 0.3 0.7 0.9 5.4 8.2 11.3 3.0 3.2 2.3 6.5 3.3 58 43.3 Carabidae 8.3 4.8 3.3 3.9 1.2 2.0 0.3 1.0 2.1 4.5 3.4 6.5 2.9 59 44.0 Calosoma scrutator – adult 0 0 0 0.4 9.7 2.0 2.3 0.5 0 0 0 0 2.4 27 20.1 Calosoma scrutator – larvae 0 0 0 0 11.4 0.5 0.8 0 0 0 0 0 2.3 18 13.4 Elateridae – adult 0.4 0.7 1.2 0.8 0.8 1.9 1.5 0.5 0.1 0.5 0.9 2.0 0.9 54 40.3 Elateridae – larvae 10.2 7.3 10.2 6.5 2.7 2.3 2.2 2.1 5.7 5.0 9.0 15.0 5.1 108 80.6 Buprestidae 0 0.3 0 0 0 0 0 0 0 0 0 0 0.01 1 0.07 Cicindellidae 0 0.5 0 0 0 0 0 0 0 0 0 0 0.0 1 0.07 Cerambycidae 0 0 0 0 0 0.04 0 2.3 0 0 0 0 0.2 4 29.8 Hydrophilidae 0 0 0 0 0 0.04 0 0 0 0 0 0 0.007 1 0.07 Scaphididae 0 0 0 0.07 0 0 0 0.8 0 0 0 0 0.07 2 1.5 Cantheridae 0 0 0 0 0 0 0 0 0 0 0.3 0 0.01 1 0.07 Cantherid larvae 0 0 0 0 0.04 0 0 0.1 0 0 0 0 0.01 2 1.5 Staphylinidae 0.3 0.3 0.2 0 1.1 0.09 0 0.5 0.1 0 0.1 0 0.3 18 13.4 Tenebrionidae 0.3 0.2 0 0 0 0.7 0 2.2 3.1 2.8 0.4 0 0.8 31 23.1 Dytiscidae 0 0 0.2 0 0 0 0 0 0 0 0 0 0.007 1 0.07 Chrysomelid larvae 0 0 0 0 0.04 0.09 0 0 0 0 0 0 0.02 2 1.5 Curculionidae 0.3 0 0 0 0 0.09 0 0.2 0.1 0 0 0.5 0.08 8 6.0 Unidentified Coleoptera adults 1.8 1.7 6.2 5.3 7.8 2.2 2.0 1.9 1.3 0 7.6 0.5 3.7 63 47.0 Unidentified Coleoptera larvae 1.5 2.5 3.2 1.3 0.3 1.6 0.2 0.7 0.3 0 0 0 0.9 21 15.7 Coleoptera pupae 0.4 0 0.5 0 0 0 0 0 0 0 0 0 0.06 2 1.5 Hemiptera (0.2) Thyreocoridae 0 0 0.2 0 0 0.04 0 0.5 0.2 0 0 0 0.07 6 4.5 Pentatomidae 0 0 0 0 0 0.2 0 0 0.07 0 0 0 0.04 2 1.5 Cynidae 0.3 0 0.3 0 0 0 0 0 0 0 0 0 0.04 4 3.0 500 Southeastern Naturalist Vol. 11, No. 3 Percent volume Frequency Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Total Percent Pyrochroidae 0.3 0 0 0 0 0 0 0 0 0 0 0 0.02 1 0.07 Coreidae 0.3 0 0 0 0 0 0 0 0 0 0 0 0.02 1 0.07 Anasa tristis 0 0 0 0 0 0 0 0 0.07 0 0 0 0.007 1 0.07 Lygaeidae 0 0.3 0 0 0 0 0 0 0 0 0 0 0.01 1 0.07 Aphidoidea – green 0 0 0 0 0 0.04 0 0 0 0 0 0 0.007 1 0.07 Membracidae 0 0 0 0 0 0.04 0 0 0 0 0 0 0.007 1 0.07 Nabidae 0 0 0 0 0 0 0.2 0 0 0 0 0 0.007 1 0.07 Unidentified Hemiptera 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0.07 Homoptera (0.2) Cicada 0 0 0 0 0 0 0 0.9 0 0.8 0 0 0.1 3 2.2 Cicada – large nymphal 0 0 0 0 0 0 0 0 0.5 0 0 0 0.06 1 0.07 Cicadellidae 0 0 1.0 0.07 0 0.09 0 0 0.07 0 0 0 0.08 4 3.0 Hymenoptera (19.4) Formicidae 8.0 20.8 11.2 29.1 23.8 30.9 46.0 4.4 6.2 12.8 2.6 9.5 19.3 128 95.5 Bee 0 0 0 0 0.2 0.3 0 0 0 0 0 0 0.08 3 2.2 Vespidae 0.08 0 0 0 0 0 0 0 0 0 0.6 1.0 0.05 3 2.2 Orthoptera (5.1) Gryllidae – Gryllus sp. 0.6 0 0.2 0.5 0.9 1.8 0.3 8.4 11.5 9.2 1.9 4.0 3.1 58 43.3 Gryllidae – Nemobius sp. 0 0.8 0 0 0.1 0 0 0 0 0 0 0 0.06 2 1.5 Blattellidae 0.6 0.5 0.7 0.4 0.4 0.3 0 0.3 6.2 0.7 0.3 0 1.0 32 24.0 Tettigoniidae 1.0 0 0 0 0.7 0.2 0 0 0.3 0.8 0 1.5 0.3 9 6.7 Acrididae 0 0.8 0 0.1 0 1.9 0 0 0 0 0 0 0.4 5 3.7 Rhaphidophoridae 0 0 0 0 0 0 3.7 0 0 0 0 0 0.2 3 2.2 Gryllotalpidae 0 0 0 0.3 0 0 0 0 0 0 0 0 0.03 2 1.5 Orthoptera – unidentified 0.2 0 0 0 0.08 0 0 0 0 0 0 0 0.03 4 3.0 Lepidoptera (13.2) Lepidoptera – larvae 10.6 14.3 11.5 6.5 10.7 19.9 6.2 3.6 7.0 8.2 24.1 13.5 11.5 114 85.0 Lepidoptera – pupae 0.6 0.3 0.5 2.0 1.9 1.2 0 0.1 0.5 0.3 0.3 1.0 1.0 29 22.0 Lepidoptera – adult 0.2 0 0 6.3 0 0 0 0 0 0 0 0 0.7 1 0.07 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 501 Percent volume Frequency Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Total Percent Isoptera (1.4) Isoptera – adult 0.2 10.7 7.2 3.3 0.3 0 0.8 0.6 0.07 1.0 0.3 0.5 1.4 23 17.2 Diptera (1.3) Chironomidae 0 0 0 0 0 0 0 0 0 0 0.3 0 0.01 1 0.07 Chironomidae – larvae 0 0 0 0 0 0.09 0 0.2 0.1 0 2.9 0 0.2 7 5.2 Tipulidae – larvae 0.08 0 3.4 1.3 0.2 0 0 0.05 0.1 0 0.4 0 0.4 15 11.2 Stratiomyidae – larvae 0 0 0.2 0 0 0 0 0 0 0 0 0 0.007 1 0.07 Unidentified Diptera – adult 0 0 0 0.1 0 0 0 0 0 0.2 0 0 0.02 2 1.5 Unidentified Diptera – larvae 4.3 0 6.3 0.07 0.1 0.04 0 0 0.1 0 0.1 0 0.7 12 9.0 Dermaptera (0.02) Dermaptera – adult 0 0 0 0 0 0.09 0 0.1 0 0 0 0 0.02 2 1.5 Odonata (0.2) Anisoptera – dragonfly 0 0 0 0 0 0 0 0 1.3 0 0 0 0.2 1 0.07 Unidentified insect (0.007) Naiad 0 0 0 0 0 0 0.2 0 0 0 0 0 0.007 1 0.07 Other Invertebrates (17.5) Chilopoda 8.6 6.3 6.5 6.3 6.6 1.8 9.3 5.1 5.5 8.5 15.9 14.5 6.5 105 78.4 Diplopoda 7.3 11.3 12.2 3.7 2.0 1.1 1.3 1.5 3.7 3.2 5.0 7.0 3.8 84 62.7 Narceus sp. 0 0 0.3 0.8 1.8 4.6 8.8 1.0 0.9 0 0 0 1.8 23 17.2 Sowbug 3.3 3.5 6.7 2.5 2.9 2.2 0.2 0.8 0.7 0.5 0.7 1.5 2.2 51 38.1 Aranaea: Spider 2.8 2. 3.2 3.5 1.6 1.0 0.8 0.7 2.1 1.7 4.9 4.0 2.1 79 59.0 Earthworm 1.6 0.3 0.8 0.6 0.04 0.3 0 0 0.5 2.3 0 5.0 0.6 21 15.7 Crab 0 0 0 0 0 0.09 0 0.2 1.1 0 0 0 0.2 4 3.0 Snail 0.4 0.5 0.7 0.07 0 0.2 0.2 0 0 0.2 0.3 0 0.2 17 12.7 Mites/Ticks 0 0 0 0 0.04 0.04 0 0 0 0 0 0 0.01 2 1.5 Slug 0 0 0 0 0 0 0 0 0 0.2 0 0 0.007 1 0.07 Invertebrate – unidentified 0 0 0 0.07 0.1 0.4 0 0 0 0 0 0 0.09 5 3.7 502 Southeastern Naturalist Vol. 11, No. 3 Percent volume Frequency Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Total Percent Vertebrates (1.9) Scaphiopus holbrookii 0 0 0 4.3 0 0 0 0 0 0 0 0 0.5 1 0.07 Frog – unidentified 2.3 0 0 0.4 0 0 0 0 0 0 0.3 1.0 0.3 6 4.5 Bufo terrestris 0.8 0 0 1.3 0.4 0 0 0 0 0 0 0 0.3 2 1.5 Lizard eggs 0 0 0 0.4 0.2 0.1 0 0 0.7 0 0 0 0.2 6 4.5 Anolis carolinensis 0 0 0 0 0 0.7 0 0 0 0 0 0 0.1 3 2.3 Scincella lateralis 0.8 0 0 0 0 0 0 0 0 0 0 0 0.07 2 1.5 Eumeces sp. 0.6 0 0 0 0 0 0 0 0 0 0 0 0.06 1 0.07 Unidentified lizard 0.3 2.0 0.3 0 0 0 0 0 0 0 0 0 0.1 5 3.7 Mammal 1.6 0 0 0 0 0 0 0 0 0 0 0 0.2 1 0.07 Bird 0 0.3 0 0 0.5 0 0 0 0 0 0 0 0.1 3 2.2 Plant material (11.8) Diospyros virginiana fruit 0 0 0 0 0 0 0 34.3 22.6 20.2 0 0 6.0 19 14.2 Endogonaceae (clumps) 2.3 2.7 0 0.07 0.04 3.3 0.8 3.0 2.4 0.2 1.4 4.0 1.6 32 23.9 Fungi 6.3 0.8 0 0 0 0.5 0 1.1 1.5 3.2 0 0 1.1 13 9.7 Nuts 0 0 0 0 0 0 0 0 0 0 11.4 0 0.6 1 0.07 Fruit 2.8 0 0 0 0 0.2 0.8 0 0 4.2 0 0 0.5 7 5.2 Melia azedarach 0 0 0 0 0 0 2.2 4.0 0 0 0 0 0.4 5 3.7 Misc. veg.: leaves, roots, etc. 1.1 0.8 0 0.3 1.1 0.2 0 0 0.1 0.5 0.3 0 0.4 11 8.2 Acorn, Quercus sp. 2.9 0 0 0 0 0 0 0 0 0 1.1 0 0.3 2 1.5 Serenoa repens seeds 0 0 0 0 0 0.04 0 0 3.1 0 0.3 0 0.4 5 3.7 Unidentified plant material 0.2 0 0.8 0.5 0 1.1 0.2 0.3 0 0 0 1.0 0.03 8 6.0 Black Cherry fruit 0 0 0 0 0 1.1 0 0 0 0 0 0 0.2 1 0.07 Vitaceae, Vitis/Ampelopsis sp. 0 0 0 0 0 0.2 0 1.3 0 0 0 0 0.1 3 2.2 Root clumps or dirt 0 0 0 0 0.3 0.1 0 0 0 0 0 0 0.08 2 1.5 Woven egg cases? 0 0 0 0 0 0.4 0 0 0 0 0 0 0.07 1 0.07 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 503 Appendix 2. Numbers of individuals of food items eaten by Nine-banded Armadillos on Cumberland Island, GA. Total Percent of Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec numbers individuals Coleoptera Scarabaeidae – adult 19 11 3 248 1126 65 3 35 20 11 6 1547 3.6 Scarabaeidae – larvae 8 3 3 17 124 184 136 28 15 3 6 527 1.2 Carabidae 108 51 40 69 33 12 2 3 47 21 16 10 412 1.0 Calosoma scrutator – adults 1 42 12 5 2 62 0.15 Calosoma scrutator – larvae 322 9 9 340 0.8 Elateridae – adult 4 3 13 29 40 58 26 9 3 3 9 5 202 0.5 Elateridae – larvae 429 71 265 268 229 134 52 76 143 116 206 167 2156 5.1 Buprestidae 1 1 0.002 Cicindellidae 2 2 0.005 Cerambycidae 1 14 15 0.04 Hydrophilidae 1 1 0.002 Scaphididae 5 5 0.01 Cantheridae 1 1 0.002 Cantherid larvae 1 1 2 0.005 Staphylinidae 3 1 1 124 2 8 2 1 142 0.3 Tenebrionidae 5 1 17 24 84 22 4 157 0.4 Dytiscidae 1 1 0.002 Chrysomelid larvae 1 7 8 0.02 Curculionidae 4 1 1 2 1 9 0.02 Coleoptera – unidentified adults 21 101 159 535 48 19 62 2 62 3 1012 2.4 Coleoptera – unidentified larvae 10 28 187 221 7 3 2 5 1 464 1.1 Coleoptera pupae 6 5 11 0.03 Hemiptera Thyreocoridae 1 2 5 3 11 0.03 Pentatomidae 1 1 2 0.005 Cynidae 2 2 4 0.009 Pyrochroidae 1 1 0.002 Coreidae 1 1 0.002 504 Southeastern Naturalist Vol. 11, No. 3 Total Percent of Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec numbers individuals Anasa tristis 1 1 0.002 Lygaeidae 1 1 0.002 Aphidoidea – green 3 3 0.007 Membracidae 1 1 0.002 Nabidae 1 1 0.002 Unidentified Hemiptera 1 1 0.002 Homoptera Cicada 2 1 3 0.007 Cicada – large nymphal 1 1 0.002 Cicadellidae 1 1 3 1 6 0.01 Hymenoptera Formicidae 991 530 223 4474 5051 5756 1236 1105 1757 1578 335 141 23177 54.4 Bee 2 3 5 0.01 Vespidae 2 1 3 0.007 Orthoptera Gryllidae – Gryllus sp. 4 1 3 11 15 3 42 52 13 8 5 157 0.4 Gryllidae – Nemobius sp. 6 1 7 0.02 Blattellidae 7 3 4 3 5 4 5 84 13 3 131 0.3 Tettigoniidae 2 2 1 1 1 7 0.02 Acrididae 1 1 8 10 0.02 Rhaphidophoridae 10 10 0.02 Gryllotalpidae 2 2 0.005 Orthoptera – unidentified 3 2 4 9 0.02 Lepidoptera Lepidoptera – larvae 236 66 222 81 586 767 186 41 82 44 192 36 2539 6.0 Lepidoptera – pupae 3 1 1 52 32 12 2 5 3 2 113 0.3 Lepidoptera – adult 1 337 338 0.8 Isoptera Isoptera – adult 103 232 1191 1221 165 60 40 1 7 15 3035 7.1 2012 J.O. Whitaker, Jr., C. Ruckdeschel, and L. Bakken 505 Total Percent of Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec numbers individuals Diptera Chironomidae 200 200 0.5 Chironomidae – larvae 3 1 5 100 109 0.3 Tipulidae – larvae 2 16 21 4 1 3 2 49 0.1 Stratiomyidae – larvae 3 3 0.007 Unid. Diptera – adult 2 2 4 0.009 Unid. Diptera – larvae 157 150 1 1 11 1 1 322 0.8 Dermaptera Dermaptera – adult 3 1 4 0.009 Odonata Anisoptera – dragonfly 1 1 0.002 Unidentified Insect Naiad 1 1 0.002 Other invertebrates Chilopoda 170 46 127 106 91 53 53 106 78 29 101 59 1019 2.4 Diplopoda 464 158 192 210 69 31 4 10 25 19 46 24 1252 2.9 Narceus 1 3 8 16 9 3 4 44 0.1 Sowbug 176 36 344 595 470 198 1 16 20 10 7 8 1881 4.4 Aranaea: spider 30 15 26 20 40 26 9 11 13 13 16 7 226 0.5 Earthworm 107 6 13 13 1 11 27 27 15 220 0.5 Crab 2 1 9 12 0.03 Snail 3 3 6 4 4 2 1 2 25 0.06 Mites/ticks 2 1 3 0.007 Slug 1 1 0.002 Invertebrate – unidentified 1 6 1 1 9 0.02 Vertebrates Scaphiopus holbrookii . 1 1 0.002 Frog – unidentified 2 2 2 6 0.01 Bufo terrestris 1 1 2 0.005 506 Southeastern Naturalist Vol. 11, No. 3 Total Percent of Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec numbers individuals Lizard eggs 3 11 1 5 20 0.05 Anolis carolinensis 1 1 2 0.005 Scincella lateralis 2 2 0.005 Eumeces sp. 1 1 0.002 Unidentified lizard 5 5 0.01 Bird 1 2 3 0.007 Plant material Diospyros virginiana 1 75 76 0.2 Endogonaceae (clumps) 1 21 1 82 15 6 7 133 0.3 Fungi 1 1 2 4 0.009 Nuts 39 39 0.09 Fruit 5 7 12 0.03 China berry, Melia azedarach 8 46 54 0.1 Misc. veg.: leaves, roots, etc. 1 1 2 1 1 6 0.01 Acorn, Quercus sp. 1 1 2 0.005 Serenoa repens seeds 40 10 50 0.1 Unidentified plant material 2 6 1 9 0.02 Black cherry fruit 78 78 0.2 Vitaceae Vitis/Ampelopsis sp. 54 54 0.1 Root clumps or dirt 3 1 4 0.009 Woven egg cases? 4 4 0.009 Seeds – blackberry, Rubus sp. 1 1 0.002 Total 3093 1279 3144 8189 9041 7598 1909 1876 2563 2019 1395 504 42,610 100