Penial characters have proven to be useful in the classification of snakes, providing bases for separating subfamilies, genera, and species. In the racer even the subspecies have trenchant penial characters by which they may be separated in some instances. The hemipenis is roughly cylindrical, but widest at the base ([Fig. 2]). The sulcus spermaticus is unbranched. Approximately the basal one-third of the hemipenis has a smooth surface, broken only by the sulcus spermaticus and by three greatly enlarged spines, which form hooks—one anterior, one posterior, and one dorsal. The dorsal hook is the largest of the three. Distal to the smooth part is a zone of small spines, each recurved and mounted on a fleshy tubercle. The zone of spines is poorly developed on the anterior side and is interrupted on the posterior side in the vicinity of the sulcus spermaticus but is best developed on the posterior side a short distance above and below the sulcus spermaticus. The spines are arranged in several oblique rows. Those of the proximal row are best developed, and there is rapid diminution in the size of those situated farther distally. Approximately the distal two fifths of the hemipenis forms a third zone, lacking distinct spines, but having numerous deep longitudinal grooves, alternating with lamellae which have fimbriated edges, and which fuse with each other and divide to form a reticulated pattern.

Relationships

The large genus Coluber is much in need of revision. Its many species, perhaps more than a score in all, occur in North America from southern Canada south to Guatemala, in eastern and southwestern Asia, in southern Europe, and in North Africa. All are active, slender-bodied snakes having smooth scales in few rows, and having large eyes with well developed vision. The North American species fall into two natural groups, the typical racers, and the whip snakes, often assigned to a separate genus, Masticophis (Ortenburger, 1928). The whip snakes are more specialized than the typical racers in having the eyes more enlarged, and the body form more slender and attenuate, with number of scale rows more reduced. The racers of the Old World are more diverse. Inger and Clark (1943) suggested a partitioning of the genus Coluber on the basis of the pattern by which scale rows are reduced, from the maximum number on the forebody to the minimum number at the posterior end of the body, supplemented by certain characters of the hemipenis and of the cephalic scutellation. Besides Coluber and Masticophis these authors recognized within the group the genus Platyceps with several species in southern Europe and southwestern Asia; Zamenis with several species in the same region and in North Africa, and Haemorrhois, a monotypic genus of Spain, North Africa and several Mediterranean islands. Although apparently valid in principle, this arrangement has not been generally followed.

Exclusive of those species groups whose assignment to the genus Coluber are somewhat questionable, the remaining species in the genus are: C. constrictor occurring throughout most of the United States and south along a narrow Atlantic coastal strip of Mexico to Guatemala; C. oaxacae of southern Mexico; and C. spinalis of northern China. C. oaxacae is poorly known as only a few specimens have been collected, but seemingly it is a near relative and derivative of C. constrictor, especially of that species' southernmost population. C. spinalis is much more distinct, as might be expected from its geographical remoteness. It is a slender, active snake, of olive coloration dorsally with 17 scale rows and a bright yellow, black-edged dorsal stripe and yellow ventral surface. It is relatively small (up to 755 millimeters snout-vent length) and is partial to riparian habitats but is also found in forests and in dry and barren regions (Pope, 1935:224-226). It is known to feed upon lizards.

Range

The common racer has been recorded in each of the 48 states of the mainland of the United States, also in New Brunswick, Nova Scotia, southern British Columbia, and southward through Mexico where it is limited to a narrow strip of east coast lowlands but extends as far as Guatemala. C. c. constrictor occupies the northeastern states and extends south into the Appalachian and Piedmont. C. c. priapus with its associated races paludicola, helvigularis, and anthicus has an Austroriparian distribution, occupying the Atlantic Coastal plain and the Gulf Region, and extending north in the Mississippi Valley to southern Illinois and Indiana. C. c. paludicola is localized with two disjunct populations—in the Everglades and on Cape Canaveral, Florida. C. c. helvigularis is even more restricted in range and is known only from the Appalachicola region of the Florida Panhandle and the adjacent corners of Alabama and Georgia. C. c. anthicus occupies much of central and western Louisiana and adjacent Texas. C. c. flaviventris occurs throughout the Great Plains, east in the "Prairie Peninsula" through Michigan and northern Ohio and west to the Rocky Mountains. C. c. stejnegerianus occurs from Matagorda Bay in Texas southward through eastern Mexico, with a seemingly isolated population in the Sierra del Carmen region of northern Coahulia. C. c. mormon occurs in the Pacific Coast states and Great Basin.

Actually, the range limits and the continuity of distribution within the area outlined are still poorly known. The species has not been recorded from the northern parts of Maine, Vermont, New Hampshire, Michigan, Wisconsin, or Minnesota, nor from northeastern New York. It is generally absent from southwestern desert areas. Records are particularly scarce and scattered in the Rocky Mountain states, suggesting that the distribution in this area may be discontinuous. In a large area comprising all of New Mexico and Arizona, the western half of Colorado, and the southern halves of Utah and Nevada, records are so scarce as to indicate that the species is there represented by only a few well isolated relict colonies. The type locality of mormon is "Valley of the Great Salt Lake," and there are numerous records from the northern part of Utah east of Great Salt Lake (Woodbury, 1931:75), but a record from Moab is the only one known to me from the southern half of the state. The only records from western Colorado are from three miles east of Fruita and two miles west of Grand Junction, Mesa County (Maslin, 1959:56). Apparently the only valid record from Arizona is that of Shannon (1950:59) from Eagar, Apache County, in the east-central part. Shannon also recorded the racer from Boulder Dam in extreme southern Nevada. Brattstrom (1955:152) has recorded the species from the lower Pleistocene of southeastern Arizona (Curtis Ranch), bearing out the idea that the racer has partly withdrawn from a range formerly occupied in the Southwest at a time when cooler and moister climate prevailed. Other fossil occurrences are of late Pleistocene age—Vero Beach and Seminole, Florida (Brattstrom, 1953a:245) and, doubtfully, Rancho LaBrea, California (Brattstrom, 1953b:376). The range of mormon has been mapped (Wright and Wright, 1949:134) as extending east to south-central Montana on the basis of one specimen allocated on the basis of two characters. Otherwise the range of mormon seems to be entirely west of the Continental Divide, well separated from that of flaviventris by desert and mountain barriers. The conspecificity of mormon with the other subspecies needs to be more thoroughly investigated, and geographic variation within mormon also merits study.


Geographic Variation