Description
A large squirrel that is highly variable in color, both individually and geographically. Over most of the range, the pelage is rusty yellowish with a pale yellow to orange (or light gray to dirty white) belly and a bushy tail bordered with fulvous-tipped hairs. In the Southeast, body may be sprinkled with yellow, white, and black, with the head mostly blackish except for the whitish nose and ears. In the vicinity of the Delmarva Peninsula, color may be pure steel-gray with no fulvous (Burt and Grossenheider 1964). Melanism is not uncommon. For adults, total length is 454-698 mm, tail length is 200-330 mm, and hind foot length is 51-82 mm (Hall 1981). Mass is 696-1361 g, with an average of about 800 g. There is a total of 20 teeth. The bones fluoresce bright red under longwave ultraviolet light (unique among normal adult mammals) (Flyger and Gates 1982).
The fox squirrel of the southeastern Coastal Plain of North America is the largest tree squirrel in the western hemisphere. Weigl et al. (1989) reported an average mass of 1006 g in North Carolina and 962 g for 44 specimens collected in the Carolinas and northern Georgia. Western fox squirrels tend to be smaller (600 to 900+ g).
Diagnostic Characteristics
Larger than SCIURUS CAROLINENSIS, in which adults are 383-525 mm in total length and 340-700 g; also, NIGER has four cheek teeth on each side of the upper jaw (CAROLINENSIS usually has five), and in NIGER the tail hairs generally are not tipped with white (Flyger and Gates 1982).
Habitat
Often in open mixed hardwood forest or mixed pine-hardwood associations, but has adapted well to disturbed areas, hedgerows, and city parks. Prefer savannas or open woodlands to dense forests (Flyger in Wilson and Ruff 1999). In Michigan, four favorite habitats in descending order: oak-hickory woodlands, beech-basswood-maple, scrub oak, and floodplains (Allen 1943). In the southeastern U.S., prinicipal habitat is mature open forests of longleaf pine and turkey oak with an open understory; adjacent bottomland hardwood forests may be occupied in summer and during periods of drought (see Handley 1991). Western range extensions are associated with riparian corridors of cottonwoods and fencerows of osage orange. Dens are in tree hollows (preferred) or leaf nests (especially in mild weather). Young are born in a tree cavity or leaf nest; tree hollows are preferred for rearing young. Individuals use up to about nine nests annually.
A few particular plant communities of the Coastal Plain appear to be important in maintaining fox squirrel populations there. The structure, age, diversity, and extent of these communities within the overall pine-oak forest provide an array of often subtle requirements for the success of the squirrel in the Southeast (Weigl et al. 1989).
In Florida and North Carolina, habitats are primarily longleaf pine (PINUS PALUSTRIS)-turkey oak (QUERCUS LAEVIS) sandhills characterized by large, well-spaced pines and an understory of scattered or clumped oaks (Moore 1953, Kantola 1986, Weigl et al. 1989, Cox 1990), although squirrels may also be found in other open pine stands, mixed pine-hardwood forests, and in ecotones between forest types. Habitat structure, specifically the size and spacing of pine and oak trees, appears to be more important than the actual species composition of the habitat (Taylor 1973, Hilliard 1979, Weigl et al. 1989). Only stands with large mature trees appear to supply adequate supplies of food and nesting sites. The large size of the eastern fox squirrel appears to adapt it to the openness of the pine-oak forest and the arrangements of habitats within the Coastal Plain. Large size is advantageous in handling the large cones of longleaf pine and in traveling along the ground between widely spaced trees, food sources, and blocks of habitat (Weigl et al. 1989).
Use of edge habitats by fox squirrels is repeatedly mentioned in the literature (Smith and Follmer 1972, Flyger and Smith 1980, Nixon et al. 1984, Kantola 1986, Weigl et al. 1989, Cox 1990). While fox squirrels in North Carolina Sandhills used pine-oak forest at least 80 percent of the time, there was marked seasonality in habitat use. During the winter months, fox squirrel activity was significantly higher in edge habitats and in bottomland forest, while in the summer months activity shifted from open pinelands and sand ridges toward moister lowlands (Weigl et al. 1989). Based on a one-year study of Florida fox squirrels, Kantola and Humphrey (1990) suggested that the best habitats may be the edges between longleaf pine savanna and live oak (QUERCUS VIRGINIANA) forest where the availability of oak mast may be most dependable from year to year.
Weigl et al. (1989) showed that fox squirrels chose nest sites surrounded by vegetation typical of longleaf pine-turkey oak forest (Wells and Shunk 1931, Braun 1964, Waggoner 1975, Christensen 1988). Fox squirrels nested in areas characterized by open, low diversity, mature forest with little understory and xeric conditions. This contrasted markedly with conditions typical of gray squirrel nest sites, which generally were in areas of closed-canopy, high diversity, dense understory, and generally mesic conditions.
Nests are of critical importance to the survival of fox squirrels throughout their range, and nest selection and use by eastern fox squirrels has received considerable attention (Moore 1957, Hilliard 1979, Flyger and Smith 1980, Weigl et al. 1989, Kantola and Humphrey 1990). Tree hollows are particularly important in rearing young and for protection from severe weather for all tree squirrels.
Fox squirrels in the southeastern U.S. often use hardwood cavities in deciduous forest, bottomlands, and mixed forests, despite the fact that the majority of foraging takes place in pine-dominated stands. Kantola and Humphrey (1990) also reported higher nest use in lower slope situations than on drier sandhill areas. Suitable hollows are fairly scarce in pine-oak forests, primarily owing to timber management practices, firewood cutting, and oak suppression (Weigl et al. 1989; Weigl, pers. comm.), but are more abundant in adjacent more mesic habitats.
Leaf nests are frequently used during warmer months and in the more southern portions of the squirrels' range. Use of leaf nests in North Carolina was associated with periods of mild weather and/or abundant food, and Weigl et al. (1989) found that nest selection during the warmest periods occurred most often in moister habitats and wetland edges. In Florida, tree cavity nests were rarely used even when abundant, comprising only 7.4% of all nest locations recorded (Kantola and Humphrey 1990).
Ecology
Fox squirrels are largely solitary animals across their geographic range, though they may congregate briefly during the breeding season or near concentrated food supplies (Bakken 1952, Havera and Nixon 1978, Benson 1980, Armitage and Harris 1982, Brown 1984, Nixon et al. 1984, Steele et al. 1984, Weigl et al. 1989, Kantola and Humphrey 1990, Handley 1991).
Much of the demography and ecology is determined by food availability and distribution. The low diversity, open pine-oak forests favored by fox squirrels in the southeastern U.S. are typified by food resources that tend to be of low abundance and very patchy distribution.
Population density in the Coastal Plain portion of the range is relatively low. Moore (1957) and Hilliard (1979) estimated densities of 0.33/ha and 0.26/ha, respectively. Weigl et al. (1989) reported densities averaging 0.05/ha. More recent estimates for northern Florida S. N. SHERMANI averaged 0.027-0.054/ha (Kantola and Humphrey 1990). In the southeastern U.S., fox squirrels are very scarce across even in the best quality habitats. In woodlots in the Midwest, average density typically is 1-4/ha but may reach 12/ha (Weigl et al., 1989, Koprowski 1994). An average of 15 gray squirrels per hectare was reported by Gurnell (1983). Numbers may vary annually, depending on weather, food availability, and the effects of diseases and parasites (Weigl et al. 1989).
The longest reported dispersal movement is 64.4 km (Allen 1943, Baker 1983, Caire et al. 1989, Kantola and Humphrey 1990).
Average home range sizes in the Sandhills of North Carolina (26.6 ha in males, 17.2 ha in females, mean convex polygon) (Ha 1983, Weigl et al. 1989) were similar to those reported from similar habitats in Georgia (males 26.4 ha, females 13.0 ha, MCP) (Hilliard 1979), but were smaller than those for South Carolina fox squirrels using pine plantations and hardwood "runners" (males 31.6 ha, females 19.3 ha) (Edwards 1986) and for the larger S. N. SHERMANI in longleaf pine forests of Florida (males, 42.8 ha; females, 16.7 ha, harmonic mean) (Kantola 1986). These home ranges are in contrast to the much smaller ranges (0.8 to 7.0 ha, MCP) reported for western fox squirrels (Baumgartner 1943, Bernard 1972, Adams 1976, Havera and Nixon 1978). The larger ranges of fox squirrels in the southeastern U.S. are not explained solely by their larger body size and diet, and they may result from the patchiness of resources in their habitat (Weigl et al. 1989, Kantola and Humphrey 1990). Home range overlap is extensive.
Weigl et al. (1989) described an intriguing relationship among the fox squirrel, its hypogeous fungal food source, and longleaf pine regeneration. They suggested that one or more fungi readily consumed by the squirrels demonstrate mycorrhizal associations with longleaf pine and that the squirrels may be an important dispersal agent of fungal spores. The squirrels naturally forage across fairly large areas, including recently burned areas and clearcuts, perhaps playing a major role in inoculating barren soils with a mycorrhizal agent necessary for vigorous growth of longleaf pine.
Nut burial may aid reforestation.
Reproduction
Fox squirrels breed at two seasons: winter-spring (young are born January-April) and, less frequently, summer-fall (young born July-September) (Moore 1957, Lustig and Flyger 1975, Weigl et al. 1989). Mating behavior in North Carolina appears to be prevalent from December to early February and again in late July and August, with most litters in North Carolina born in late February and March (Weigl et al. 1989).
Gestation lasts 44-45 days (Moore 1957, Kantola 1986). In the southeastern U.S., most young are born in March-April; sometimes there is a smaller second peak in July-August (Weigl et al. 1989). Litter size generally averages 2-3. Mean litter sizes of 2.3 and 2.27 (Moore 1953) have been reported for Florida fox squirrels. For the southeastern U.S., Weigl et al. (1989) reported litter sizes (mean = 2.5) that equaled or were well below the lowest litter sizes reported for other sciurids. Many authors have suggested that older females may produce a second litter in good habitat. However, Weigl et al. (1989) recorded no instance of a second litter in eight years and suggested that no records of a second litter occur in the literature on fox squirrels in the southeastern U.S. Weaning occurs during the period when the young are 8-13 weeks old. In Florida, young are dependent on their mother for about 3 months. Earliest time of first breeding usually is the second calendar year. Family group may stay together until female is ready to mate again. Maximum reproductive longevity of females is about 12-13 years.
Reproductive output, including the percentage of sexually mature individuals that reproduce, increases with food abundance (Hilliard 1979, Kantola 1986, Weigl et al. 1989). Reproduction and litter size in North Carolina were significantly associated with food availability in the previous fall-winter periods. During poor mast years, juvenile survival increased, while adult condition declined, resulting in an overall negative impact on squirrel populations (Koprowski 1991). Juvenile squirrels are particularly sensitive to food shortages due to relatively low fat stores, higher metabolic demand, low experience in locating and processing scarce food, and perhaps low social rank (Koprowski 1991).
Few data exist on fox squirrel longevity. Data from midwestern fox squirrel populations suggest longevities of 13 years in captivity and six to seven years in the wild (Flyger and Gates 1982). The large size and low recruitment rates of fox squirrels in the southeastern U.S. suggest relatively long life spans (Moore 1953, Weigl et al. 1989). Weigl et al. (1989) suggested a life span of seven years in the Sandhills of North Carolina, although no data on age or survivorship have been reported.