Forest floor components utilized by eastern box turtles include litter, natural depressions, soft soils, brush, and woody debris. Eastern box turtles often seek shelter by digging a form in moist soil or leaf litter. They sleep within forms at night and rest in them during the day. Their carapaces are partially to completely covered by soil, litter, or vegetation while in the form [71]. The average depth of forms in Arkansas was 0.21 inch (0.53 cm) below the surface [63]. Other cover, such as brush piles, woody debris, briar patches, and tangled vines is utilized throughout the day [71]. Hatchling and juvenile eastern box turtles often hide under leaf litter, which does not offer protection against fire [34]. The microhabitat in which neonate eastern box turtles were found had significantly more leaf litter (p=0.007), less herbaceous cover (p<0.001), and shorter vegetation (p<0.001) than random sites. Neonate eastern box turtles were found at microsites with high light intensity and low canopy cover, which led to higher temperatures than at nearby microsites [38].
The most important habitat features for hibernating eastern box turtles include cavities or natural depressions (such as stump holes and other hollows) filled with deep litter, as well as soft soils, thick brush, and woody debris [12,14,24,73]. Some eastern box turtles overwinter in depressions along gully bottoms and hillsides [12]. As winter gets progressively colder, eastern box turtles dig deeper into litter and soil to gain more protection from cold [12]. Snow cover helps insulate hibernating turtles [1]. In rare circumstances, eastern box turtles successfully hibernate while submerged in a stream or pond [11,44]. Eastern box turtles may also utilize burrows dug by other wildlife [34]. Juveniles that hatch late in the season may overwinter in the nest [52]. Multiple eastern box turtles are occasionally found overwintering in the same location [12].
Fire mortality for the eastern box turtle can be highly variable. For instance, after a prescribed fire in a tallgrass prairie habitat in Missouri, 22 eastern box turtles were found alive and 20 were dead [39]. Babbitt and Babbitt [3] discovered 17 % mortality in an eastern box turtle population in Florida after what was probably a wildfire. The study site was located near the Everglades where prefire vegetation was characterized by "thick undergrowth" [3]. In Oklahoma, 25 eastern box turtles and ornate box turtles (T. ornata) were found dead after a fire, while only 3 box turtles (Terrapene spp.) were found alive [6]. Allard [2] suggested that spring fires may be detrimental to eastern box turtles when they become active after hibernation. This statement was based on carcasses discovered in burned areas at this time of year, although data on eastern box turtles killed in such fires were not given. No eastern box turtles were captured in a prescribed burn site in Maryland. However, captures were very low in the cut-over site, white oak-willow oak-red maple (Quercus alba-Q. phellos-Acer rubrum) forest, and loblolly pine-mixed hardwood forest [54]. The low capture success in this study limits the inferences that can be drawn in regards to the effects of fire on eastern box turtles.
Ernst and others [34] suggested that eastern box turtles occupying burrows likely escape fire completely. However, some eastern box turtles that died in an apparent wildfire in Florida were found in burrows [3]. Hatchling and juvenile eastern box turtles appear to hide under litter, which exposes them to fire, rather than burrowing or creating forms[34].
Eastern box turtles appear incapable of escaping advancing fires, so they are frequently found with burn scars [3,13,34]. Many eastern box turtles that survive fire while in their forms are badly burned, often with extensive damage to the shell [3,34]. Eastern box turtles can regenerate part to all of damaged or burned shells [64,68]. The ability of eastern box turtles to regenerate their shells after being burned is possibly an adaptation for survival in fire-prone environments [68].
In general, eastern box turtles will eat anything edible that fits into their mouths [23]. Examples include fungi, snails and slugs (Gastropoda), insects (Insecta), spiders (Arachnida), centipedes (Chilopoda), millipedes (Diplopoda), earthworms (Annelida), carrion, and vegetation [10,23,48,71,74]. Fungi may comprise 10% to 55% of the eastern box turtle diet [10,71,74]. In Kentucky, snails and slugs comprised and average of 52.5% of the diet by volume [10]. Consuming carrion appears common and includes reptile, amphibian, mammal, bird, and fish carcasses [48]. Specific examples include eastern racers (Coluber constrictor), Fowler's toads (Bufo fowleri), eastern ribbonsnakes (Thamnophis sauritus), and red-bellied snakes (Storeria occipitomaculata) [48]. An eastern box turtle was once seen feeding on a cow (Bos taurus) carcass [13]. Eastern box turtles have been observed eating dead toads (Bufo spp.) [58]. An eastern box turtle in Oklahoma was observed preying upon plains leopard frog (Rana blairi) tadpoles in a dry pond [7]. Dead rats (Rattus spp.) are eaten by captive eastern box turtles [1].
Plant materials eaten by eastern box turtles include leaves, berries, roots, flower buds, and seeds [10,23,48,71,74]. Eastern box turtles have been observed eating half-flower (Scaevola taccaca) berries, cactus (Cactaceae) fruits, and seagrape (Coccoloba uvifera) in Florida [23]. Eastern box turtles consume and disperse seeds of pond-apple (Annona glabra), Florida silver palm (Coccothrinax argentata), fig (Ficus spp.), redgal, (Morinda umbellata), sapodilla (Manilkara zapoda), crowngrass (Paspalum spp.), mangroveberry (Psidium longipes), Everglades greenbrier (Smilax coriacea), Key thatch palm (Thrinax morrisii), saw palmetto (Serenoa repens), and Long Key locustberry (Byrsonima lucida) [51]. They may also consume and disperser seeds of jack-in-the-pulpit (Arisaema spp.), mayapple (Podophyllum petatum), black cherry (Prunus serotina), summer grape (Vitis aestivalis), common elderberry (Sambucus nigra ssp. canadensis), black huckleberry (Gaylussacia baccata), blue ridge blueberry (Vaccinium vacillans), white mulberry (Morus alba), American pokeweed (Phytolacca americana), Indian strawberry (Duchesnea indica), Virginia strawberry (Fragaria virginiana), blackberry (Rubus spp.), muscadine grape (V. rotundifolia), and frost grape (V. vulpina) [8]. Other vegetative foods of eastern box turtles include fox grape (V. labrusca), cherry (Prunus spp.), pear (Pyrus spp.,) sweetroot (Osmorhiza spp.), American wintergreen (Pyrola americana), groundcherry (Physalis spp.), grasses, and mosses [8,48,71,74].
Data collected on the effects of fire on eastern box turtle habitat are limited. Frequent fires may limit the distribution or population size of eastern box turtles in some areas. This may be especially true in Florida [13]. Schwartz and Schwartz [65] determined that spring fires may burn off leaf litter covering hibernating eastern box turtles, exposing them to freezing temperatures. Since eastern box turtles show a preference for forested and other woody habitats (see Preferred Habitat), fires that reduce or eliminate forested habitats could be detrimental to eastern box turtle populations. However, eastern box turtles are found in successional habitats [54,60,65], which indicates they could adjust to changing landscapes that are caused by fire.
FIRE REGIMES: The following table provides fire return intervals for plant communities and ecosystems where the eastern box turtle is important. Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find FIRE REGIMES".
Community or ecosystem Dominant species Fire return interval range (years) maple-beech Acer-Fagus spp. 684-1,385 [16,77] silver maple-American elm Acer saccharinum-Ulmus americana <5 to 200 sugar maple Acer saccharum >1,000 [77] bluestem prairie Andropogon gerardii var. gerardii-Schizachyrium scoparium <10 [49,59] bluestem-Sacahuista prairie Andropogon littoralis-Spartina spartinae <10 [59] birch Betula spp. 80-230 [75] sugarberry-America elm-green ash Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica <35 to 200 Atlantic white-cedar Chamaecyparis thyoides 35 to >200 beech-sugar maple Fagus spp.-Acer saccharum >1,000 black ash Fraxinus nigra 77] green ash Fraxinus pennsylvanica <35 to >300 [33,77] cedar glades Juniperus virginiana 3-22 [42,59] yellow-poplar Liriodendron tulipifera <35 shortleaf pine Pinus echinata 2-15 shortleaf pine-oak Pinus echinata-Quercus spp. <10 slash pine Pinus elliottii 3-8 slash pine-hardwood Pinus elliottii-variable <35 sand pine Pinus elliottii var. elliottii 25-45 [77] South Florida slash pine Pinus elliottii var. densa 1-15 [57,69,77] longleaf-slash pine Pinus palustris-P. elliottii 1-4 [57,77] longleaf pine-scrub oak Pinus palustris-Quercus spp. 6-10 Table Mountain pine Pinus pungens <35 to 200 [77] pitch pine Pinus rigida 6-25 [9,45] pocosin Pinus serotina 3-8 pond pine Pinus serotina 3-8 [77] eastern white pine Pinus strobus 35-200 [75,77] eastern white pine-northern red oak-red maple Pinus strobus-Quercus rubra-Acer rubrum 35-200 loblolly pine Pinus taeda 3-8 loblolly-shortleaf pine Pinus taeda-P. echinata 10 to <35 Virginia pine Pinus virginiana 10 to <35 Virginia pine-oak Pinus virginiana-Quercus spp. 10 to <35 sycamore-sweetgum-American elm Platanus occidentalis-Liquidambar styraciflua-Ulmus americana <35 to 200 [77] eastern cottonwood Populus deltoides <35 to 200 [59] quaking aspen-paper birch Populus tremuloides-Betula papyrifera 35-200 [32,77] black cherry-sugar maple Prunus serotina-Acer saccharum >1,000 oak-hickory Quercus-Carya spp. <35 northeastern oak-pine Quercus-Pinus spp. 10 to <35 [77] oak-gum-cypress Quercus-Nyssa spp.-Taxodium distichum 35 to >200 [57] southeastern oak-pine Quercus-Pinus spp. <10 white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra <35 northern pin oak Quercus ellipsoidalis <35 bear oak Quercus ilicifolia <35 bur oak Quercus macrocarpa <10 [77] oak savanna Quercus macrocarpa/Andropogon gerardii-Schizachyrium scoparium 2-14 [59,77] chestnut oak Quercus prinus 3-8 [77] northern red oak Quercus rubra 10 to <35 [77] post oak-blackjack oak Quercus stellata-Q. marilandica <10 black oak Quercus velutina <35 live oak Quercus virginiana 10 to<100 [77] cabbage palmetto-slash pine Sabal palmetto-Pinus elliottii <10 [57,77] blackland prairie Schizachyrium scoparium-Nassella leucotricha <10 Fayette prairie Schizachyrium scoparium-Buchloe dactyloides <10 [77] eastern hemlock-yellow birch Tsuga canadensis-Betula alleghaniensis 100-240 [75,77]Common mammalian predators of box turtles are northern raccoons (Procyon lotor), skunks (Mephitis mephitis and Spilogale spp.), American minks (Mustela vison), coyotes (Canis latrans), domestic and feral dogs (Canis familiaris), and rats (Rattus spp.) [2,35]. Other potential predators include American badgers (Taxidea taxis), Virginia opossums (Didelphis virginiana), foxes (Vulpes and Urocyon spp.), nine-banded armadillos (Dasypus novemcinctus), and weasels (Mustela spp.) [35]. Birds that prey upon eastern box turtles include American crows (Corvus brachyrhynchos), Mississippi kites (Ictinia mississippiensis), barn owls (Tyto alba), herring gulls (Larus argentatus), and turkey vultures (Cathartes aura) [35,46].
The shells of young box turtles are not strongly ossified until they reach several years of age, making them vulnerable to predators [26]. Hatchling box turtles may fall prey to shrews, birds, eastern chipmunks (Tamias striatus), bullfrogs (Rana catesbeiana), and snakes [4,44,53]. Copperheads (Agkistrodon contortrix), cottonmouths (Agkistrodon piscivorus), eastern racers (Coluber constrictor), and other snakes may swallow young eastern box turtles whole [35,47,56].
Eastern box turtle eggs are preyed upon by snakes such as scarletsnakes (Cemophora coccinea), hog-nosed snakes (Heterodon spp.), common kingsnakes (Lampropeltis getula), pinesnakes (Pituophis melanoleucus), and eastern ratsnakes (Elaphe obsoleta), as well as ants and other invertebrates [2,35,58].
Eastern box turtles show a preference for forests, especially bottomland forests and edge habitats [6,12,17,19,31,52,54,60,63,65,70,71]. Eastern box turtles in Mississippi inhabited longleaf pine-slash pine (Pinus palustris-P. elliottii) forests ranging from early to mature to late successional stands [60]. Mixed stand habitats in Maryland, dominated by loblolly pine (P. taeda) that originated as agricultural land, were undergoing succession to an oak-maple (Quercus-Acer spp.) forest [54]. In Missouri, eastern box turtles occupied a previously cultivated ridge [65]. In an Oklahoma study, an eastern box turtle habitat was partially dominated by range and pasture [6]. Eastern box turtles occasionally inhabit pastures and marshes in Kansas [17]. Eastern box turtles utilized grasslands, open lawns, and meadows in Arkansas and Florida [28,63]. Conversely, eastern box turtles seemed reluctant to use grassy, herbaceous, and low brush-covered fields in New York [52]. In another New York study and in Maryland, eastern box turtles did not appear to discriminate between habitats because they were found in bottomland hardwood and mixed pine-hardwood forests, shrublands, mixed grasslands, wetlands, riparian zones, croplands, and rural developed areas [19,54,70,71,72]. However, abundance of eastern box turtles in each of these habitats was not reported.
High humidity seems to be one of the most important factors in habitat selection. Mean relative humidity in eastern box turtle habitat in Arkansas was above 80%. Average total ground cover around forms (shallow depressions dug by eastern box turtles) was 40.27%, while the litter averaged 54.34% cover and a depth of 1.41 inches (3.57 cm). Average grass cover at the forms was 12.16%, forb cover was 14.32%, and shrub cover averaged 8.81%. Average canopy cover above forms was 56.05% with an average canopy height of 36.65 feet (11.17 m) [63]. The undergrowth in a Maryland forest was littered with heaps of woody debris, fallen branches, logs, and stumps [72].
Forests provide cool areas and high humidity during the heat of the summer [63]. The most preferred forest habitats were those with the most moisture and highest diversity [52]. In Mississippi, eastern box turtles were found in habitats characterized by gently rolling hills dissected by intermittent and perennial streams [60]. Box turtles (Terrapene spp.) seem to avoid ridges where moisture is low and generally avoid steep hillsides and embankments [24]. Eastern box turtles may use virtually any habitat during rainy weather [13], and they are most active after rain showers [58].
In addition to humid environments, eastern box turtles utilize open water extensively. Eastern box turtles swim across streams and other bodies of water [71,76]. Eastern box turtles are also known to spend hours or days soaking in puddles, lakes, streams, and wet gullies [1,24,71]. In Tennessee, they utilized temporary ponds during periods of high temperature and low precipitation [31]. Eastern box turtles, especially juveniles, may dry out and perish during long periods of drought [1]. Neonates may congregate in open water or seek shade after hatching to avoid dehydration and heat stress [1,11].
Elevation: In a review, Dodd [27] notes that eastern box turtles in New England are common from sea level up to 490 feet (150 m) in elevation, and rare to 705 feet (215 m). In the southern Appalachians, eastern box turtles are common from sea level to 4,300 feet (1,300 m) but rare at higher elevations [27]. Wilson and Friddle [80] noted that eastern box turtles are common in valleys below 1,000 feet (300 m) in elevation, but are rare on ridges above 2,000 feet (600 m) in West Virginia.
Density/Home Range: Average densities of eastern box turtle populations can vary widely and may reflect differences in habitat quality and other environmental factors. For instance, in Indiana, density estimates were 2.7 to 5.7 eastern box turtles/ha [78]. A density estimate in a Virginia population was considerably higher at 35 eastern box turtles/ha [79]. In Missouri, eastern box turtles had an average density of 7.3 to 10.9/acre (18.1-27.0/ha) [66]. In Maryland, an average of 4.1 to 5.9/acre was found (1.7-2.4/ha) [43,71]. In Tennessee, there were 12.3 eastern box turtles/acre (5.0/ha) on average [30]. An eastern box turtle population in Florida had an estimated density of 14.9 to 16.3 adults/ha [25,62].
Eastern box turtles do not appear to be territorial because they are commonly found grouped together under cover or in close proximity to each other. Eastern box turtles occupy the same home range year after year. However, females may leave their home ranges to lay eggs [71]. Home ranges of eastern box turtles in Missouri averaged 3.6 acres (1.5 ha) for females and 3.8 acres (1.5 ha) for males [65]. Average home ranges over a 19-year period in Missouri were 12.7 acres (5.1 ha) for females and 12.9 acres (5.2 ha) for males at the same location [66]. Home ranges in New York may average 4.35 to 17.20 acres (1.76-6.96 ha) [52]. Home range size of eastern box turtles in Virginia averaged 19.5 acres (7.9 ha) [79]. On average, home ranges of eastern box turtles ranged between 4.65 acres (1.88 ha) and 5.58 acres (2.26 ha) in Tennessee [31].
Sexual maturity for the eastern box turtle is reached at 5 to 10 years of age [55]. Dodd [25] noted in a review that eastern box turtles may live to exceed 100 years on occasion, while reaching 50 to 60 years may be fairly common. Most wild eastern box turtles likely only live up to 5 years [25].
Hibernation: Eastern box turtles are active from April to November in northern parts of their range, as well as on warm winter days [1,11,27,52,63,65]. Hibernation begins October to November, and emergence from hibernation begins in March [12]. Eastern box turtles do not appear to hibernate in Florida. Several environmental cues have been identified relating to the timing of hibernation. Generally, eastern box turtles hibernate between the last severe autumn frost and the 1st spring frost [35]. One study concluded that they begin emerging from hibernation when ambient temperatures reach 65 °F (18 °C) in spring [17]. Another study suggested that box turtles (T. carolina and T. ornata) emerge from hibernation when subsurface soil temperatures (4-8 inches (10-20 cm) below the soil surface) are at least 45 °F (7 °C) for a minimum of 5 days [41].
Reproduction: Mating occurs May to October in Missouri [65] and has been observed in late November in Florida [22]. Nesting occurs May to July [27,36], with hatching from August to November [27,36,44]. Clutches may contain 1 to 9 eggs with an average of 3.67 to 5 eggs per clutch being typical [13,27,46]. Multiple clutches of eggs may be laid in a single year [79]. Eastern box turtle nests are roughly as deep as the female can reach with her hind legs, approximately 2 to 4 inches (6-10 cm). Eggs are laid primarily during rainy and overcast weather [18]. Incubation lasts 60 to103 days [17,18,36]. Males primarily develop at cooler temperatures while females predominantly develop at higher temperatures [44].
Fire at any time of year appears to be harmful to eastern box turtles since they are unable to escape [3,13,34]. Fire mortality has not been studied extensively, but mortality is consistently high in studies that have examined the effects of fire on eastern box turtle populations [3,6,39]. Given the high mortality rate, frequent fires may severely reduce an eastern box turtle population [13]. However, given that at least a few individuals appear to survive fire, a turtle population may be able to recover if the site has a long fire return interval and the forested habitat has not been greatly reduced.
High-severity fires that kill trees and scorch canopies would likely be detrimental to eastern box turtles since they favor forests [6,12,17,19,31,52,54,60,63,65,70,71]. The removal of the litter layer by fire could also be detrimental because litter is used extensively for cover throughout the year [12,38,63]. The adverse affects of removing litter from the forest floor early in the year would probably be short-term if the leaves in the canopy fell later in the year. However, an autumn fire occurring after most leaves have fallen would have more severe effects on eastern box turtles since a deep litter layer is crucial during hibernation [12,14,24,73]. Timing of fire may be less of a problem in Florida since eastern box turtles typically do not hibernate in that location [35]. More research is needed to address these possibilities.
Research on the effects of fire on eastern box turtle populations and habitat is lacking. Concern for the eastern box turtle already exists over populations that have been isolated through habitat fragmentation [21]. More research is needed to determine if fire in these habitat fragments would be detrimental to the isolated eastern box turtle populations.
