The Reptiles of South Carolina: Foreword by J. Whitfield Gibbons

Introduction

Reptiles are vertebrate animals that are ectothermic amniotes. The former class Reptilia is paraphyletic and does not reflect evolutionary relationships among amniotes (reptiles, birds, and mammals). Crocodilians are archosaurs, which are more closely related to birds and dinosaurs than to lizards, snakes, or turtles. Snakes and lizards are lepidosaurs, which are more closely related to the tuatara, a primitive New Zealand reptile, than to turtles or crocodilians. Using DNA sequencing, scientists are uncovering the phylogeny of living vertebrates and adjusting the taxonomy accordingly.

Taxonomy is classification of organisms and should reflect evolutionary relationships. Traditionally external morphology such as color patterns or scale counts were used to classify reptiles. With recent advances in sequencing and analyzing DNA, the scientific community is witnessing significant changes to animal taxonomy. I used the recent classification for species in this book. Where changes are so new that they are not present in field guides, I included the older name as well. This is evident in the taxonomy of species with large geographic ranges and in classification at the genus and family levels.

Taxonomists give binomial names to species using the method developed by Linnaeus over 200 years ago. The name includes the genus name, which is capitalized, and the specific epithet, which is not capitalized. Scientific names are in Latin or Greek and allow scientists to communicate across the globe. English common names are easier for the lay public but vary across space and time and often reflect misinformation or mythology. Standard English common names for reptiles I have used are in Crother et al. (2012).

Species with broad geographic ranges often exhibit geographic variation in morphology. When these differences adhere to portions of a species range they have been described as subspecies. Subspecies are defined as an aggregate of evolutionarily similar populations of a species that inhabit a geographic subdivision of the range and differ from other populations of that species (Mayr and Ashlock, 1991). Subspecies are given a trinomial name like Cemophora coccinea copei for the northern scarlet snake. Wide-ranging reptiles exhibit variability in color pattern, and subspecies have been described based on this variation. Where different subspecies meet, they interbreed, and a narrow to wide zone of intergradation exists. Populations of animals in the intergradation zone exhibit phenotypes that are intermediate between the named subspecies. Recent studies of phylogeography of many species have determined subspecies do not reflect evolutionary history. I did not recognize subspecies where genetics has shown them to not be evolutionarily relevant.

FIG. 1. County map of South Carolina

Organization of Species Accounts

The species accounts have been organized so a morphological description of a typical adult is given, followed by a discussion of sexual dimorphism and ontogenetic variation exhibited. A description of the geographic range and habitat characteristics follows. Range maps show where the species has been found in South Carolina. Solid symbols indicate the locality has a voucher specimen in a scientific collection from that site. Open circles are based on literature records and credible sightings or photographs. Some symbols represent more than one locality when the localities are close. The section on taxonomy and geographic variation provides information on subspecies occurring in the state and recent name changes. An overview of the ecology and life history is given in the natural history section. This information is based on studies done in South Carolina whenever possible. The section on conservation status addresses conservation issues at the state level and where appropriate on the national or global level (Gibbons, Scott, et al., 2000). I reviewed the scientific literature for South Carolina populations of each species in the last section.

A Brief History of Herpetology in South Carolina

South Carolina has a rich history in zoology and natural history that parallels the state’s history. While this section is not meant to be exhaustive it points out major contributions. An early work depicting amphibians and reptiles from South Carolina was Natural History of Carolina, Florida and the Bahama Islands published in the period 1731–43 by the English naturalist Mark Catesby (1683–1749). This series was published in two volumes with illustrations of 23 species of reptiles that occur in the Carolinas. Catesby lived in Charleston from 1722 to 1725, and many of his specimens went to the British Museum of Natural History in London (Sanders and Anderson, 1999). The American bullfrog (Lithobates catesbeiana) is named in his honor.

Another important eighteenth-century naturalist was William Bartram (1739–1823), who wrote Travels through North and South Carolina, Georgia, East and West Florida, which was first published in 1791 (Adler, 1979). Bartram’s work resulted from four years in the Southeast collecting seeds and plant specimens for English horticulturalists and botanists. His illustration of American alligators (Alligator mississippiensis) is his most famous drawing, but his description is inaccurate (Adler, 1979). He illustrated soft-shelled turtles (Apalone sp.), discussed rattlesnakes, the moccasin (presumably Agkistrodon piscivorus), pigmy rattlesnakes (Sistrurus miliarius), rough green snakes (Opheodrys aestivus), ribbon snakes (Thamnophis sauritus), chicken snakes (presumably Pantherophis alleghaniensis), pine snakes (Pituophis melanoleucus), green lizards (Anolis carolinensis), and several species of tortoises (freshwater turtles).

Nineteenth-century physician John E. Holbrook (1796–1871) was born in Beaufort, South Carolina, and grew up in Wrentham, Massachusetts (Worthington and Worthington, 1976). Holbrook received his A.B. degree from Brown University in 1815 and his M.D. degree from the University of Pennsylvania in 1818. He had a private medical practice in Charleston in 1822. Like many physicians of his day Holbrook had an interest in natural history and in ectothermic vertebrates. His North American Herpetology, whose second edition was published in five volumes in 1842, was an attempt to describe and illustrate all species of amphibians and reptiles occurring in what was then the United States. Almost 150 different species, by the taxonomy of the day, were included in this ambitious work. Holbrook began this project during the 1820s and collected specimens in all east coast states except Florida. Five salamanders, four anurans, four turtles, eight snakes, and the genera Scaphiopus, Crotaphytus, and Pituophis, which he described, are still valid taxa (Sanders and Anderson, 1999). North American Herpetology was well received by Holbrook’s peers on both sides of the Atlantic and earned him the title of father of American herpetology.

For almost 250 years the Charleston Museum (ChM) has been an important institution for herpetological research. Founded in 1773 by the Library Society of Charleston, it is the oldest natural history museum in the United States (Sanders and Anderson, 1999). Edward Burnham Chamberlain (1895–1986), ChM curator of zoology from 1924 to 1952, began bringing bird specimens to the museum with his twin brother, Rhett, in 1905 (Sanders and Anderson, 1999). Chamberlain began publishing on South Carolina herpetology in 1928, writing about salamanders, frogs, pigmy rattlesnakes (Sistrurus miliarius), alligators, birds, and small mammals from 1910 to 1939. He published at least eight papers on South Carolina herpetology. The salamander Eurycea chamberlaini is named in his honor. Albert Schwartz (1923–92), ChM curator from 1952 to 1956, published four papers on South Carolina herpetology. An avid collector, he increased the size of the vertebrate collections at ChM with South Carolina specimens. Albert E. Sanders, curator of natural sciences at ChM from 1968 to 2012, published The Reptiles of Columbia, South Carolina and Vicinity in 1966, with a revised edition in 2004.

Julian D. Corrington (1887–1979) was a biology professor at the University of South Carolina from 1921 to 1926. He discovered the black swamp snake (Seminatrix pygaea) in South Carolina (Corrington, 1927) and published a herpetological inventory of the Columbia area (Corrington, 1929). Unfortunately Corrington’s specimens were lost (Sanders and Anderson, 1999). Julian R. Harrison III (1934–2009) grew up working with the ChM and became a professor at the College of Charleston in 1963. He published a list of coastal zone amphibian species (Harrison, 1978) and described Eurycea chamberlaini from South Carolina (Harrison and Guttman, 2003).

Herpetological studies in the piedmont lagged behind those of the coastal plain. Clemson University scientists worked in the upstate and mountains. Andrew L. Pickens (1890–1969) published separate lists of amphibians and of reptiles in 1927. He was a professor at Furman University until his defense of evolution led to his dismissal in 1926 (Sanders and Anderson, 1999). Franklin W. Sherman (1877–1969), an entomologist at Clemson University, prepared lists of piedmont herpetofauna and shared them with E. B. Chamberlain at the ChM. Richard Montanucci was a herpetologist at Clemson University from the 1970s to about 2010 and studied amphibians in the upstate (Montanucci, 2006).

The largest contributor to herpetology in South Carolina has been the Savannah River Ecology Laboratory (SREL). The Savannah River Site (hereafter SRS) was established in 1951 for the enrichment of radioactive isotopes. The SRS encompasses approximately 780 km² along the Savannah River in parts of Aiken, Barnwell, and Allendale Counties. It is found within the upper coastal plain including some fall line sandhills (Gibbons and Semlitsch, 1991). The first herpetological surveys were conducted by Harry W. Freeman of the University of South Carolina (Freeman, 1955a; 1955b). Although an ichthyologist, Freeman had an interest in herpetology and was hired at the University of South Carolina in 1951 to survey fishes, amphibians, and reptiles of the SRS. The University of Georgia’s SREL was opened in 1962 on the SRS (Cothran et al., 1991). The herpetology laboratory, started in 1967 and made major contributions to herpetology, especially ecology of amphibians, turtles, and alligators. Over one million specimens of 100 species of amphibians and reptiles have been marked on the SRS by researchers at the SREL herpetology lab (srelherp.uga.edu). J. Whitfield Gibbons, his students, and staff have published dozens of papers on the ecology of the herpetofauna on the SRS. Gibbons (1978) is a species list of reptiles inhabiting the state’s coastal zone. Justin D. Congdon and his students have also published extensively in turtle ecology and other aspects of herpetology. I. Lehr Brisbin Jr. and colleagues have conducted long-term ecological studies of alligators and radioecology studies of many vertebrate species on the SRS. As of this writing Kurt A. Buhlmann and Tracey D. Tuberville run the SREL herpetology lab.

Although no treatise covering the herpetology of the entire state has been published, an annotated list by James T. Penney (1900–1964) was published in 1952. The list covered all species of herpetofauna known to occur in South Carolina at that time. The popular work Amphibians and Reptiles of the Carolinas and Virginia (Martof et al., 1980) included color photographs and coarse-scale range maps of all species known in the region and was revised by Beane, Braswell, et al. (2010).

Physiographic Regions of South Carolina

South Carolina ranks 40th of the 50 states in size with around 78.9 million hectares. The state contains three physiographic regions with the coastal plain being the largest, followed by the piedmont and the Blue Ridge, the smallest. Each will be treated separately in the following section. Many species of reptiles are restricted to one or two of these physiographic regions, whereas others show morphological differences from one region to another. The fall zone is the boundary between the coastal plain and piedmont. It is where rivers plunge off the piedmont onto the coastal plain, and rapids occur in rivers of this region. The fall zone can be up to 2.4 km wide (C. Murphy, 1995).

FIG. 2. Physiographic regions of South Carolina. The fall line sandhills are part of the upper coastal plain adjacent to the fall line.

Atlantic Coastal Plain Physiographic Region

The Atlantic coastal plain extends from southern New England to Florida (Rohde et al., 2009) and westward as the Gulf of Mexico coastal plain. Geologically it consists of unconsolidated sand and clay sediments with little exposed rock. The coastal plain encompasses all South Carolina southeast of the fall zone, which is about two-thirds of the state. The fall zone runs northeast to southwest from near Cheraw, Chesterfield County, through Columbia to North Augusta, Aiken County (C. Murphy, 1995; Rhodes, 1998). Coastal plain topography consists of flattened terraces gradually becoming gently rolling hills near the fall zone. Elevations range from sea level to 220 m in the fall line sandhills, and it is 190–240 km wide (C. Murphy, 1995). The coastal plain is divided into the lower coastal plain closest to the coast and the upper coastal plain contacting the fall zone. The Orangeburg Scarp is a wave-cut ridge running diagonally across the state at elevations of 55–66 m (C. Murphy, 1995) and is the boundary between these two parts of the coastal plain (Porcher and Rayner, 2001). The coastal plain contains the highest number of reptile species in South Carolina. Several species of reptiles such as spotted turtles, chicken turtles, Florida cooters, eastern glass lizards, banded water snakes, and glossy crayfish snakes are restricted to the coastal plain.

The lower coastal plain includes seven terraces and six escarpments formed by changing sea levels during the Pliocene, Pleistocene, and Holocene epochs (C. Murphy, 1995). This region has the lowest elevation, ranging from sea level to 84 m. Owing to the flat nature of the landscape it the has the most wetlands. Some reptiles such as the island and mimic glass lizards and the eastern diamond-backed rattlesnake are restricted to the lower coastal plain. The portion of the lower coastal plain directly influenced by tides or salt spray is the coastal zone or Maritime Strand extending 16 km inland and includes maritime forests, tidal creeks, and salt marshes (Porcher and Rayner, 2001). The diamond-backed terrapin is the only reptile limited to the coastal zone in South Carolina.

South Carolina has 295 km of coastline, which is divided into three regions. The Grand Strand is natural beaches extending from Winyah Bay in Georgetown County to North Carolina (C. Murphy, 1995). The central 29 km of coastline between Winyah Bay and the Santee delta includes the Cape Romain National Wildlife Refuge. The beaches on Cape Romain are important loggerhead turtle nesting grounds. The southern portion of the coast, the Sea Island Complex, extends for 160 km from the Santee delta to the Savannah River (C. Murphy, 1995) and is composed of numerous barrier islands, which are ringed in salt marsh and support maritime forest.

The upper coastal plain lies between the fall zone and the Orangeburg Scarp with elevations of 84–150 m (Beane, Braswell et al., 2010). The topography is more rolling than the lower coastal plain and is well drained with fewer wetlands. The upper coastal plain closest to the fall zone contains the fall line sandhills, which form a discontinuous band across the state 8–48 km wide (C. Murphy, 1995). These deep Cretaceous to Miocene sands extend from Chesterfield County across Kershaw, Richland, Lexington, and Aiken Counties. The best remnant longleaf pine forests occur in the fall line sandhills. Although no reptiles are restricted to the upper coastal plain, many are very abundant in the fall line sandhills, including the rare southern hog-nosed snake and pine snake.

Piedmont Physiographic Region

The piedmont is that part of South Carolina that lies northwest of the fall zone and extends to the base of the Blue Ridge Escarpment. The piedmont physiographic province extends from southern New York to Alabama (Rohde, et al., 2009). This region is 175 km wide and constitutes one-third of South Carolina. Piedmont topography is rolling hills closer to the mountains, which give way to gently sloping terrain closer to the coastal plain. Igneous and metamorphic rocks underlie the piedmont (C. Murphy, 1995). Soils of the piedmont are high in clay and reddish in color. Elevations in the South Carolina piedmont range from 122 m near the fall zone to 365 m adjacent to the Blue Ridge Escarpment (Rohde, et al., 2009). The piedmont also has monadnocks or rocky outcrops such as Kings Mountain in York County and Paris Mountain in Greenville County (Porcher and Rayner, 2001). Owing to the dissected nature of the landscape few natural wetlands are present in the piedmont. No reptiles are unique to the South Carolina piedmont, but many widespread North America species are found there but not on the coastal plain, including painted turtles, northern water snakes, and queen snakes.

Blue Ridge Physiographic Region

The Blue Ridge Mountains are the easternmost ridges of the Appalachian Mountains. They range from Virginia to northern Georgia. Only 2% of South Carolina lies within these mountains, which occur in Greenville, Oconee, and Pickens Counties. On the east, the mountains are defined by the steep Blue Ridge Escarpment, which rises about 615 m above the piedmont. These mountains are made of granites and gneisses (Porcher and Rayner, 2001). This is the coolest and wettest part of the state, receiving more than 180 cm of precipitation each year. Sassafras Mountain straddles the state line with North Carolina and is the highest point in South Carolina, with an elevation of 1,085 m (Rohde et al., 2009). Few reptiles are restricted to this part of the state, but those that are include the rare bog turtle and the coal skink.

Major Habitats of South Carolina

Owing to a wide variety of natural habitats and a relatively mild climate South Carolina has a rich reptile fauna. Statewide South Carolina averages about 123 cm of rain yearly, with more than 180 cm in the Blue Ridge Mountains and 100 cm in the midlands (Rohde et al., 2009). Average temperatures vary from 17°C along the coast to 12°C in the mountains. At the time of European settlement, South Carolina was predominately forested.

Forest Types

South Carolina still retains much forest, but the character and species composition have changed over the past three centuries (Ware et al., 1993; Earley, 2004). Most of the coastal plain, not in wetlands, was covered by longleaf pine (Pinus palustris) forests, which had open understories with a diverse herbaceous layer. Today these forests occupy less than 3% of their former range (Ware et al., 1993) and are considered a rare ecosystem. Longleaf pine is fire adapted, and the forests it dominated are fire climax communities. Post–Civil War logging removed these trees, and fire suppression prevented their regeneration. Many herbaceous understory plants are also fire adapted, such as wiregrass, which requires growing season fires to reproduce (Earley, 2004). Fires burned frequently in the coastal plain and were started by lightening or Native Americans. Owing to the flat topography fires burned for miles before encountering wetlands, which stopped them.

Longleaf grows in dense stands or scattered in savannas. The understory composition and tree density depend on the slope, aspect, soil type, water table, and fire regime. The understory is dominated by turkey oak (Quercus laevis) and grasses. In flatwoods the understory harbors saw palmetto and gallberry. Bushes like Vaccinium, legumes, and composites are important understory plants (Means, 1996).

Longleaf pine forests are important because of their rarity and high biodiversity. The canopy allows enough light to reach the ground for a diverse herbaceous plant community to exist. Plant species richness ranges from 150 to 300 species per hectare and was measured as high as 42 species per 0.25 m² (Means, 1996). Breeding bird diversity is higher in these forests than in other southeastern forests, and herpetofaunal species richness exceeds 100 in the South Carolina coastal plain. The herbaceous understory and sandy soils make excellent habitat for many herpetofaunal species. At some sites xeric sandy ridges are right next to swamps. Going from a desert-like habitat to a swamp covers only a few meters in distance and a two-meter change in elevation; yet there is almost complete floral and faunal turnover. Rare reptiles dependent on this habitat include the gopher tortoise (Gopherus polyphemus), eastern diamond-backed rattlesnake (Crotalus adamanteus), and pine snake (Pituophis melanoleucas).

Longleaf pine (Pinus palustris) flatwoods at Hobcaw Barony, Georgetown County, with drift fence for sampling amphibians and reptiles.

In the absence of fire other trees outcompeted longleaf pine; thus the southern mixed hardwood forest (SMHF) replaced the piney woods (Skeen et al., 1993). This replacement forest consists of loblolly pine, sweetgum, beech, oaks, hickories, and magnolias. Trees grow in dense stands, and little light reaches the ground. There are few herbaceous plants and associated insects relative to the longleaf pine forests. Today SMHF is more widespread than longleaf pine and has much lower biodiversity.

Areas with extensive sand deposits are xeric and referred to as sandhills. They occur along the fall zone or along rivers. The alluvial sandhills support the same vegetation as fall line sandhills, which is open longleaf pine forest or savanna with an understory of turkey oak, wiregrass, prickly pear cacti, and yucca. Although no reptiles are restricted to sandhills, many are abundant, including rare species like gopher tortoises, southern hog-nosed snakes, and the pine snakes.

The piedmont was settled early, and European invaders intensively farmed this area, denuding it of topsoil and vegetation. By the late eighteenth century, settlement of the Carolina piedmont was at its peak, and by 1830 settlement was complete (Skeen et al., 1993). Little information exists concerning the natural vegetation of the piedmont. Skeen et al. (1993) contend it was an oak-hickory-yellow poplar forest with an open understory due to fire. Pines were a small component and were limited to disturbed sites or sandy soils. Today piedmont forest is primarily oak-hickory-pine forest with loblolly and shortleaf pine.

Sandhills with turkey oak (Quercus laevis) and longleaf pine (Pinus palustris), Carolina Sandhills National Wildlife Refuge, Chesterfield County.

On flat coastal plain sites with poor drainage so the water table is close to the ground’s surface, flatwoods occur (Earley, 2004). Some flood periodically, whereas others are drier. They are dominated by longleaf pine with pond pine or slash pine present. Flatwoods that burn frequently have carnivorous plants in the understory. Fire-tolerant shrubs like wax myrtle, saw palmetto, hollies, and fetterbush may be present. Eastern diamond-backed rattlesnakes are fond of flatwoods.

Along rivers and larger creeks in the piedmont and the coastal plain riparian or bottomland hardwood forests occur (J. Nelson, 1986). They have well-developed canopies and include red maple, sweetgum, loblolly pine, yellow poplar, oaks, sycamore, ash, and elms. These forests drain poorly and flood during much of the year. Tree species richness is higher in this forest type than other habitats in South Carolina. Congaree National Park in Richland County is an example of primary floodplain forest. Common five-lined skinks (Plestiodon fasciatus) inhabit this forest type.

Maritime forest occurs on barrier islands and behind coastal dunes (J. Nelson, 1986). Trees are subject to excessive wind and salt spray. It contains live oak, palmetto, southern magnolia, slash pine, loblolly pine, or longleaf pine. Maritime forests burn infrequently but are vulnerable to disturbances from the ocean or smothering by dunes. Hunting Island State Park is an example of maritime forest. Eastern coral snakes (Micurus fulvius) frequent this type of forest.

Piedmont forest, Pickens County.

Riparian or bottomland forest, Francis Marion National Forest, Charleston County.

Maritime forest, Hunting Island State Park, Beaufort County.

Appalachian hardwood forest, Table Rock State Park, Pickens County.

Mountain bog, Greenville County.

The Blue Ridge Mountains in South Carolina were covered in Appalachian oak forest, which is now dominated by white and chestnut oaks with hemlock, birch, and hickory (Stephenson et al., 1993). They were dominated by American chestnuts, which were the most abundant tree in the southern Blue Ridge prior to European invasion (Stephenson et al., 1993). Chestnuts died out early last century from an introduced fungus. This habitat can be found in Caesar’s Head State Park in Greenville County. The eastern milk snake (Lampropeltis triangulum) is found in this habitat. Mountain bogs are sphagnum moss–dominated wetlands of the southern Appalachians with vegetation that is a combination of southern endemic and northern relictual species (Richardson and Gibbons, 1993). In South Carolina they are restricted to the mountains, and the bog turtle (Glyptemys muhlenbergii) inhabits mountain bogs.

Wetlands and Aquatic Habitats

Four main river drainages occur in South Carolina. The Pee Dee drainage encompasses the northern coastal plain and adjacent piedmont. The Santee drainage takes in most of the piedmont and some coastal plain south of the Pee Dee drainage. The Edisto River drains most of the southern coastal plain. The Savannah River drainage includes all southern and western portions of the state. Water flows faster on the piedmont and slows after crossing the fall zone onto the coastal plain. Oxbow lakes form in floodplains of major rivers as they cross the coastal plain. Black water rivers have clear, darkly stained water and originate on the coastal plain. They include the Black, Little Pee Dee, and Edisto Rivers. Reptiles like river cooters (Pseudemys concinna concinna), brown water snakes (Nerodia taxispilota), and alligators inhabit the state’s rivers and streams.

Swamp, Lynches River County Park, Florence County.

Swamps are closely associated with rivers and streams on the coastal plain. The dominant trees are water tupelo and cypress. Swamps develop on the coastal plain where there is little topography and rivers lack discreet boundaries. Along larger rivers, swamps form a band between the bottomland hardwoods and the river channel. Swamp forest is associated with lentic habitats like Carolina bays, ponds, and lakes. Swamps provide habitat for reptiles like the common musk turtle (Sternotherus odoratus), cottonmouth (Agkistrodon piscivorus), and alligators.

Carolina bays are shallow elliptical depressions on the coastal plain oriented with their long axes in a northwest to southeast direction. Bays range from less than a hectare to several thousand hectares in size, and larger bays are several kilometers long. They are named for the bay forest growing in them with sweet bay, red bay, and loblolly bay the major components. Carolina bays have either clay or peat bottoms, and many are bordered by sand rims, supporting sandhills communities of pines and turkey oak (C. Murphy, 1995). Many exhibit vegetative zonation, with up to five vegetation zones, from the drier outer edges to open water in the center (Richardson and Gibbons, 1993). Most are ephemeral wetlands, with only 10% supporting fish populations. Although no reptiles are endemic to bays, they have diverse assemblage of them. Turtles reach high biomass in Carolina bays (Congdon and Gibbons, 1989). Woods Bay State Natural Area in Clarendon and Sumter Counties is a large permanent Carolina bay.

Alligators, Caimans, Crocodiles, and Gavials

Order Crocodilia

There are about 25 species of crocodilians in three families that occur throughout the tropical regions of the world. Crocodilians are archosaurs and more closely related to birds and dinosaurs than they are to turtles or squamates. Crocodilians have a four-chambered heart, a secondary palate in the roof of the mouth, extensive parental care, and they make vocalizations. Two crocodilians are native to the United States and include the American crocodile (Crocodylus acutus), restricted to southern Florida, and the American alligator (Alligator mississippiensis), which is found from Texas to North Carolina. Crocodiles occur worldwide in the tropics, and gavials (family Gavialidae) are two species in southern Asia with extremely slender snouts.

Family Alligatoridae: Alligators and Caimans

These broad-snouted crocodilians are found in the Western Hemisphere and in eastern China. Only the Chinese alligator (Alligator sinensis) occurs outside the Western Hemisphere. Caimans include seven species of neotropical alligatorids. The American (A. mississippiensis) and Chinese alligators are the only temperate zone crocodilians. Alligators and caimans can be distinguished from crocodiles by the fact that when their mouths are closed no lower jaw teeth are visible. Owing to their much narrower jaws, some teeth on the lower jaw are visible when a crocodile’s jaws are closed.

American Alligator (Alligator mississippiensis)

DESCRIPTION: The American alligator is a large, broad-snouted crocodilian with a laterally compressed tail. The eyes and nostrils occupy an elevated position on the skull, allowing them to penetrate the surface of the water while the animal is submerged. The eyes have vertical pupils and a reddish shine at night when illuminated. Several rows of bony plates known as osteoderms protect the animal’s back. The webbed feet have five toes on the front limbs but only four toes on the hind feet. There are two keels that run along the edges of the dorsal surface of the tail, which fuse into a single keel about half way down the tail. Adults are dark gray to black in color without any conspicuous markings, whereas the hatchlings are crisply banded in black and yellow. In juveniles, the banding begins to disappear with some light regions remaining along the sides and on the lower jaw. Unlike lizards, which have a transverse cloacal slit between the hind limbs, the cloacal opening of crocodilians is longitudinal.

American alligator (Alligator mississippiensis), Clarendon County. Photograph by G. W. Long.

Males are larger than females. Females rarely exceed 3 m in length, whereas males can exceed 5 m in length and weigh up to 300 kg. The most reliable way to determine the sex of alligators is to probe the cloaca, where the penis of the male is readily apparent. Hatchlings are about 25 cm in length and weigh approximately 40 to 60 g. Alligators could not easily be confused with any other reptile native to South Carolina.

DISTRIBUTION AND HABITAT: The American alligator occurs from Florida to North Carolina and west to Texas. In South Carolina alligators occur in appropriate habitat throughout the coastal plain. Rhodes (1998) showed six piedmont localities, and Platt et al. (1999) discussed their occurrence in Lake Hartwell in Oconee County. These are not established populations and represent introductions or dispersal events by sub-adult animals.

FIG. 12. Geographic distribution of the American alligator (Alligator mississippiensis) in South Carolina. Closed circles are localities supported by voucher specimens, and open circles are observations and literature records.

Alligators prefer aquatic habitats with still or slowly moving water and gently sloping banks. They are found in rivers, large streams, swamps, marshes, lakes, impoundments, canals, sloughs, and brackish tidal wetlands. In South Carolina alligators are more numerous along the coast and seem to have benefitted from the construction and maintenance of former rice fields and other impoundments.

TAXONOMY AND GEOGRAPHIC VARIATION: The American alligator is monotypic; therefore no subspecies are recognized.

NATURAL HISTORY: Alligators in South Carolina mate in the spring, most often May and June. Courtship begins with males making bellowing vocalizations with their heads held out of the water. These low-pitched advertisement calls proclaim ownership of their territory and attract females. The female enters the male’s territory exhibiting submissive postures, and she may bellow in response to the male’s vocalization, if she is interested in mating. Copulation occurs underwater in the male’s territory.

Crocodilians, like most species of turtles, exhibit temperature-dependent sex determination. This means that the sex of the offspring is determined by the temperature to which the developing embryo is exposed (Rhodes and Lang, 1996). Specifically higher and lower temperatures during the middle trimester of development produces females, and males are produced by intermediate temperatures. In South Carolina female alligators lay eggs in mid-June, which hatch in late August or early September (Rhodes and Lang, 1995; 1996). South Carolina alligator nests exhibit an incubation period of 62 to 66 days (Roe et al., 2004). Nests average just over 4 m (range 0–45 m) from the water’s edge. The nests are mounds of living or dead vegetation that are assembled and guarded by the female. Clutch sizes of alligator nests in South Carolina average about 45 eggs and range from 22 to 57 eggs (Rhodes and Lang, 1995; 1996). Females normally guard the nest, and the hatchlings will vocalize at hatching so the mother can help them leave the nest and enter the water. Females may remain with their young for more than two years.

Hatchling alligators in South Carolina weigh from 41 to 63 g and are 23 to 26 cm in length (Brisbin, Gaines, et al., 1996; Roe et al., 2004). Males grow about 20 cm in length per year for their first six years, whereas females grow at 18 cm per year until they are six years old (P. Wilkinson and Rhodes, 1997). After age six growth rates slow to just over 7 cm per year for males and just over 6 cm per year for females. They reported that females reached sexual maturity at about 6.5 years old at about 2.1 m in length. The largest females found in this coastal population were 2.94 m in length, and the largest male was 3.84 m long (P. Wilkinson and Rhodes, 1997). Further inland on the SRS, T. Murphy (1981) reported that alligators do not reach maturity until they are 10 years old.

Alligators are generalist carnivores and take a wide variety of vertebrate prey. There are few published accounts of alligator feeding habits in South Carolina. Chamberlain (1930) reported that a specimen from Charleston County contained mullet remains and metal leg bands of little blue and tricolored herons. In North Carolina alligators have been known to feed on insects, spiders, millipedes, crayfish, blue crabs, numerous species of fishes, mudsnakes, water snakes, turtles, and several species of aquatic birds such as herons, ducks, and geese (Palmer and Braswell, 1995). Juvenile specimens consumed most of the invertebrates. They