Lowlands to mountains; desert, prairie, brushland, woodland, open coniferous forest, farmland, marshes. Eastern populations occupy sandy pine woods and dry shaly ridges; midwestern population inhabit prairies; western and Mexican populations occur from coastal grasslands and forests through deserts into montane forests (Sweet and Parker 1990). Terrestrial, fossorial, and arboreal. Remains underground in cold weather and during hot midday period in summer (Fitch 1956, Zappalorti et al. 1983, Burger et al. 1988); may occupy mammal burrow (Schroder 1950, Fitch 1958) or dig own burrow (pine snakes and bullsnakes, aided by a pointed snout and enlarged rostral scale, are accomplished burrowers in loose soil [Franz 1992]; Carpenter [1982] estimated that burrowing PITUOPHIS could move up to 3400 cubic cm of soil in an hour). Eggs are deposited in burrows excavated by the female in loose soil (Moore 1893; Zappalorti et al. 1983; Burger and Zappalorti 1986, 1991), spaces beneath large rocks or logs, or possibly small mammal burrows (Ernst and Barbour 1989, Franz 1992).

Good quality pine snake habitat appears to be characterized by xeric, pine-dominated or pine-oak (50 to 80% pine) woodland with an open, low understory established on sandy soils. Longleaf pine sandhills appear to represent critical habitat over much of the southeastern U.S. Pine snakes also require forest openings, with level, well-drained sandy soils and little shrub cover, as nesting and hibernation sites.

New Jersey: habitat is restricted to the open Pine Barrens of the southern portion of the state (Zappalorti et al. 1983). Ten radio-telemetered individuals tended to be found in dense vegetation that provided shade and moist, cool ground cover (Burger and Zappalorti 1988). Snakes were inactive 41% of the time, basked 36%, were moving 20%, and nested 3%. Selected habitat was primarily (93% of locations) pitch pine-scrub oak (QUERCUS MARILANDICA) uplands (50-80% pine); Atlantic white cedar (CHAMAECYPARIS THYOIDES) swamps and lowland pine areas were avoided. Snakes spent roughly equal amounts of time in human-altered and undisturbed habitats, with no major seasonal habitat shifts other than at nesting and hibernation. Early successional habitats in sites that had been previously disturbed by human activity appeared to be of crucial importance in both foraging and nesting (Zappalorti and Burger 1985). Habitat selection of radio-telemetered pine snakes differed appreciably from the apparent habitat selection of hand-captured or randomly encountered snakes (Burger and Zappalorti 1988). Thermoregulation appeared to be important in habitat selection. Snakes were inactive at very high and very low temperatures, when they remained under wet leaves or inside hollow logs. On windy days they sought cover in blueberry (VACCINIUM spp.) patches and basked partially covered by vegetation, possibly to avoid dehydration (Burger and Zappalorti 1989).

Hibernacula and summer dens in New Jersey usually were beside old fallen logs, with entrance tunnels following decaying roots into soil. Snakes in hibernacula were at depths of 50-110 cm. Hibernacula were characterized by lower surface and soil temperature, higher surrounding leaf cover (mean = 55%), were closer to trees (mean = 130 cm), and had more trees within 10 m (mean = 138) than random points in clearings. In general, hibernacula were relatively more protected sites than random points and were less vulnerable to daily temperature fluctuations. Hibernacula appeared to have been excavated by the snakes themselves. Typically they had an average of eight side chambers and 642 cm of tunnels. Except for hatchlings, most snakes in hibernacula were located in individual chambers off the main tunnel and not in groups. All snakes were found at depths of 50 to 111 cm (mean = 79 cm). All hibernacula were within 100 m of nesting sites and most were only 10 to 15 m from nest chambers. See Burger et al. (1988) for further details. In New Jersey, eggs were deposited in nests as much as 25-30 cm deep. Nest sites often were re-used in subsequent years (Burger et al. 1988; Burger and Zappalorti 1991, 1992). Excavation took 2-4 days to complete and not all females that excavate nests were are gravid (Burger and Zappalorti 1991). Of 93 nest sites studied by Burger and Zappalorti (1986, 1991) all were located in large clearings with less than 10% tree cover in pitch pine-scrub oak xeric uplands. Nest sites were in soft sand in open, unvegetated sections of clearings, either recently cleared or burned areas with exposed sand. Clearings used for nesting averaged 166 m long and 79.2 m wide with slopes less than 14 degrees. Ground cover averaged less than 5% over the entire nest, less over the mouth and egg chamber. Nests were excavated in soft packed sand with little or no humus. Snakes appeared to select a particular range of sand hardness that provided ease of digging but was firm enough to avoid burrow collapse. Soil moisture may have been important in site selection (Burger and Zappalorti 1986, 1991). Nests ranged between 90-305 cm (mean = 187.6 cm) in length, averaged about 14 cm wide, and eggs were found at an average depth of about 21 cm below the surface. All nests were located in clearings or along roads or railroad beds. Clearings were usually man-made (only 1 or 20 clearings appeared to be naturally created by fire). Burger and Zappalorti (1986) suggested that man-made clearings were crucial to nesting ecology in the New Jersey Pine Barrens and may have improved nesting habitat for this population.

North Carolina: longleaf pine (PINUS PALUSTRIS) sandhills (Brimley 1944).

South Carolina: Gibbons et al. (1991) described the habitat as sandy old fields and turkey oak (QUERCUS LAEVIS)-pine forests.

Louisiana: both P. M. LODINGI and P. M. RUTHVENI are restricted to longleaf pine forests and second growth longleaf pine- blackjack oak (QUERCUS MARILANDICA) associations (Fugler 1955, Walker 1965). Both P. M. LODINGI and P. M. RUTHVENI appear to use the burrows of pocket gophers, gopher tortoises, and armadillos (Dundee and Rossman 1989). Texas: Mitchell and Tinkle (1960) reported observing P. M. RUTHVENI foraging in a seasonally dry, acid bog, where the vegetation consisted of loblolly pine (PINUS TAEDA) and shortleaf pine (PINUS ECHINATA) with a dense sweetbay (MAGNOLIA VIRGINIANA) and cyrilla (CYRILLA RACEMIFLORA) understory. Florida: subspecies MUGITUS is found in xeric sites, occurring primarily in longleaf pine-turkey oak woodlands, but also in sand pine (PINUS CLAUSA) scrub, pine flatwoods on well-drained soils, and old fields on former sandhill sites (Franz 1992). Pine snakes seek out open habitats surrounded by wetlands during drought (Franz 1992). Franz (1992) also reported Florida pine snakes to actively seek burrows of gopher tortoises (GOPHERUS POLYPHEMUS) as shelter.

NATURE SERVE STATE HABITAT COMMENTS:

On Coastal Plain these snakes are found in relatively open forests of longleaf pine, wiregrass and turkey oaks; and also in dry piney flatwoods. Little is known of mountainous habitats.

REPRODUCTIVE HABITAT COMMENTS: