Part 8

Many observers have published accounts of the courtship and/or mating of the racer, but all of these are, to some degree, incomplete. Because of the widely different circumstances, and the different viewpoints of the observers involved, the several accounts give much different impressions of sexual behavior in this species. Either singly or combined, the published accounts do not provide an adequate description of the process.

My own observations, made both under natural conditions and in large outdoor enclosures, are likewise somewhat incomplete, but indicate that the whole sequence of courtship and mating is divisible into the following well-defined stages: 1) the finding of a receptive female by the male; 2) the persistent following of the female by the male, who courts her by lying extended along her body and performing writhing movements, with periodic interruptions during which he momentarily leaves the female and courses rapidly through the grass around her; 3) the acceptance of the male by the female, signalled by the raising of her tail and the almost instantaneous intromission; 4) the dragging of the passive male by the female while he is firmly attached to her during the period of coitus; 5) separation of the pair and involution of the male's hemipenis.

Even in the breeding season, racers that were confined in enclosures usually were either indifferent to each other or responded with reactions of fear or hostility. In moving they tended to follow the edges, and often two moving in opposite directions would approach each other; when this occurred, one snake might strike at the other with a short jab that seemed to be mostly bluff, and then would dart away. The males, being smaller, were usually the more wary.

Sexual behavior was noticed on only a few occasions. Several large adult males were less wary than others and usually manifested curiosity or interest toward other racers. My most complete observations of sexual behavior were made on May 18, 1962, when a newly caught adult male was added to an enclosure of 100-foot circumference already containing several racers, two of which were large adult females. Within half an hour the male was found courting one of the females. She was lying in a loose coil, with the male extended along her. At my approach the female darted away in alarm for approximately three feet, and the male moved with her, so swiftly and adroitly that he maintained contact and was in approximately his original position with respect to the female when she stopped.

Spasmodic rippling movements passed down the body of the male as he lay in contact with the female. These movements lasted several seconds, increasing in intensity, alternating with longer periods of little or no movement. As each period of vigorous writhing reached its climax, the male's head jerked forward and backward several times in seeming excitement. The female's behavior was mostly passive. She seemed to be receptive, but from time to time, without any noticeable warning, she darted away for several feet as she had when the pair was first discovered. Each time the male darted forward with her, maintaining contact while she moved. These swift movements of the female seemed to be spontaneous, at least in most instances there was no evident cause for alarm. The female's movements seemed to stimulate the male's interest rather than to discourage him. In most instances the female moved only four to five feet, then stopped abruptly or turned back. She would stop in a loose resting coil, in thick grass, with the male lying over her. Often she coiled in such a way that the posterior end of her body was beneath her forebody, but this did not seem to deter the male from moving the posterior end of his body into position beside hers. After a sudden change in the female's position, the rear of the male's body would perform groping movements along that of the female until his cloacal region was approximately opposite hers. The male sometimes had his chin pressed against the female's back, especially when he was moving forward along her, but more often his head was raised, and frequently was as much as 18 inches from the female's head.

At intervals averaging approximately ten minutes, during a little more than an hour of observation, the male would suddenly dart away from the female, and with unusually rapid and animated movements, he would move around her in an irregular and devious course, sometimes as far as five feet away, but usually within 18 inches. Usually on each such expedition several or many circuits were made; then the male would return to the female and would glide rapidly along her until he attained the mating position. A period of especially vigorous courting movements would follow.

At 12:55 p. m. it was necessary for me to discontinue observations, and I left the female confined in a cloth bag. Returning at 1:20 p. m. I found that the male was not displaying interest in the female confined in the bag, nor in the other female loose in the enclosure. The first female was released from the bag, and was out of sight for approximately four minutes. When relocated she was again attended by the male, who was carrying on courtship even more vigorously than he had before. At 1:35 p. m. the male achieved intromission. Although the pair was under observation at the time intromission occurred, the actual eversion of the hemipenis was not seen because the snakes were partly concealed by dense vegetation. There was a sudden flurry of movement, the male's head waving and his body thrashing. In an instant these violent movements subsided, and after a few seconds the female began to crawl forward slowly. The male had relaxed, and relinquished his contact with the female anteriorly. As she moved away he was dragged after her tail-first. He made slight backward wriggling movements that perhaps aided in maintaining sexual contact. The female's restlessness increased, and in eight minutes she dragged the male in a circuitous course a distance estimated to be between 20 and 30 feet. At 1:40 p. m. the pair was ten feet from the point where copulation had begun. The female showed increasing inclination to climb, raising her head and forebody against the trunks of saplings, and finally reaching up one to a branch 20 inches above the ground, and climbing first along the branch and then farther up the main trunk. As she progressed the male was lifted from the ground, dangling limply suspended by his hemipenis and its base had become exposed. At 1:43 p. m. separation occurred and the male dropped into the grass. Semen dripped from the cloacae of both snakes. That from the female was tinged with blood. The individuals involved in this observation were kept in the enclosure subsequently but no further sexual behavior was noted.

Contrary to the popular belief that these racers have permanent mates, all available evidence indicates that they are promiscuous, and two or more males may simultaneously court the same female in the brief spring breeding season. On May 24, 1960, while I was walking in a hilltop field of brome grass, a sudden movement attracted my attention to three racers lying alongside each other. Only the posterior parts of their bodies and their tails were visible. Two were males and were performing the characteristic slow writhing movements against the body of the female from either side. Although the heads were not in view, the snakes may have been able to see me through the screening vegetation; after I had watched for approximately 20 seconds, all three suddenly took alarm, for no apparent cause, and scattered.

Further evidence of promiscuity is provided by the account of Ellicott (1880:207) who wrote regarding the eastern subspecies: "I noticed a ball of black snakes (Bascanion constrictor L) rolling slowly down a steep and stony hillside ... about two miles above Union Factory, Baltimore County, Md. ... kept together by procreative impulses." It was stated that this observation was made in early spring. "Snake balls" have often been observed, and described in the literature; usually the snakes involved were garter snakes (Thamnophis) or water snakes (Natrix). Seemingly, typical aggregations consist of a single adult female and several or many males attempting to mate with her. There is a distinct possibility that the snakes involved in Ellicott's observations were misidentified.

Sexual behavior of the racer is in most respects remarkably similar to that of the common garter snake, Thamnophis sirtalis, well known through the work of Blanchard and Blanchard (1942). In studying sexual behavior of racers, several observers have failed to differentiate between the different stages of the mating process, and have assumed that copulation was occurring when actually only the precopulatory behavior was observed. In an early description of courtship in this racer in Kansas, Brons (1882:365) stated that the female "at times, seems to toy with the male, indisposed to yield to his importunities, though pressed with ardor. To avoid his suit, at times, she will dart through grass, among stones, or enter a crevice. Should he be able to reach his mate while within a hole, he is not slow in bringing her to the surface, again to be repulsed. Upon an unbroken ground the sexual union is less prolonged. Here she is unable to free herself from his quick and effectively directed moves. In case she attempts to quit him, a coil is thrown about her body, and his head laid flat upon her neck, and replaced as promptly as dislodged, evidently in the endeavor to propitiate her."

Another account probably based on courtship rather than copulation is that of Wright and Wright (1957:135), who described the behavior of a pair of C. c. priapus on Billy Island, Okefinokee Swamp in southern Georgia, on May 8, 1921, as follows: "They were stretched out, more or less coiled ... the rear parts of the bodies from the vent were entwined. The female, or smaller one seemed to have its tail around that of the male. There were contortions or quiverings from time to time.... May 8, 1921: Jackson Lee saw black snakes entwined, the male seizing the female by the top of the neck."

Blanchard and Blanchard (op. cit.) have described the dragging of the male by the female during coitus in the garter snake, and the temporarily inseparable bond formed between members of a pair by the recurved spines of the engorged hemipenis, but it has not been generally recognized that the process is much the same in other colubrines. Cottam (1937:229) described and photographed mating in a pair of C. c. mormon in Utah. The copulating racers were shown in a loose coil lying alongside each other with tails intertwined. However, when disturbed by the observers, these racers made frantic efforts to escape, crawling in a spiral course, while remaining attached and intertwined, "with no evident attempt to separate" during approximately a quarter hour of observation.

The racer is notorious for its aggressive behavior and occasional alleged attacks on humans in the breeding season. The tendency has doubtless been much exaggerated, especially in the verbal second- or third-hand accounts based on the alleged observations of eye-witnesses. Nevertheless, the supposition that large adults will sometimes pursue or attack humans when disturbed is well substantiated. In most of the instances known to me, it is the large eastern subspecies, C. c. constrictor, involved in these incidents, and seemingly the smaller racers of the Middle West, far West and South are less inclined to behave aggressively. In May 1958 two pairs of large racers were confined in a semicircular wire enclosure thirty feet across and open on top, and with natural vegetation, at the Reservation headquarters. Often in approaching the cage I saw two or more racers in close association, but because of sheltering vegetation, and the snakes' timidity observation was difficult. On May 19 a pair were lying partly extended in loose coils, but immediately the female took alarm and darted away, breaking loose from the male; his hemipenis was exposed, and underwent involution and retraction in approximately 30 seconds. Unlike the female, the male on this occasion did not attempt to escape, but turned to face me with a show of aggressiveness. Probably copulation was in its final stages when the disturbance occurred.

Circling of the female racer by the male from time to time in the course of courtship has not been recognized by previous observers as a part of the mating pattern, but Pope (1944:171) described somewhat analogous behavior, probably modified by unnatural conditions of captivity and the crowding of many racers in one cage. Pope, citing earlier observations by Noble, wrote: "When sexually excited, the male blacksnakes dash wildly about before paying court to individual females. In captivity these dashes excite all specimens confined together. A male, after picking out a mate, moves his chin lightly along her back, while undulations run forward along his sides and he extends his tongue now and then. Later he throws the part of his body near his vent over the corresponding part of the female, the two tails sometimes becoming loosely intertwined."

Recorded dates of mating for the species are all in spring, but indicate a span of many weeks for the breeding season, and this spread results in part from geographical differences. Published records are as follows:

Subspecies constrictor May 12, 1930, in Ohio (Conant, 1938:55)

Subspecies priapus May 8, 1921, in Georgia (Wright and Wright, 1957:135) May 9, 1921, in Georgia (Wright and Wright, 1957:135)

Subspecies flaviventris May 3, 1931 (two pairs) in Missouri (Boyer and Heinze, 1934:195) April 18, 1936, in Missouri (Anderson, 1942:210) May 12, 1928, in Kansas (Gloyd, 1928:123)

Subspecies mormon June 10, 1927, in Utah (Cottam, 1937:229) July 7, 1938, in California (Cunningham, 1959:17)

In the course of my live-trapping, I occasionally found more than one racer in a trap. As might be expected from the low yield per trap, such double or multiple captures were relatively rare. Chance, and unusually strategic placement of certain traps were doubtless contributing factors. May and October, being the most productive months for trapping, yielded a high proportion of these combined captures. Some involved an adult and an immature snake, or two adults of the same sex. Eliminating all these, there remain 44 heterosexual captures of adults. These latter captures are significantly concentrated in their seasonal distribution and indicate a spring breeding season; 34 were in May, six were in June and four were in October. Eight of the May records and one June record each involved a trio of snakes--two males and a female in every instance. Distribution of the spring records, grouped in five-day intervals, was as follows:

May 11 to May 15: 14 June 5 to June 9: 0 May 16 to May 20: 5 June 10 to June 14: 2 May 21 to May 25: 8 June 15 to June 19: 2 May 26 to May 30: 13 June 20 to June 24: 0 May 31 to June 4: 1 June 25 to June 30: 1

Approximately 87 per cent of the records fell in the twenty-day interval, May 11 to 30, which is regarded as the main breeding season. Presumably males continue to be at the peak of breeding condition and continue to search for females after the latter have become unreceptive, partly explaining the scattering of records through most of June.

Several of the females found in traps with males in May had abundant active sperm in their cloacae and oviducts and probably had been inseminated within a few hours of the time they were checked. Others lacked sperm, but the cramped quarters inside the traps may have effectively prevented the consummation of courtship, especially when two males were confined with the same female. None of the females trapped with males in October was found to be inseminated, and it seems doubtful whether copulation ever occurs at that time of year, although males have motile sperm and seem to be in breeding condition then.

Cycle of the Male

Cloacal smears indicate that males mature sexually and first produce sperm in August and September when they are a little more than a year old. Insofar as could be determined, there was no sexual activity at this time of year, and actual breeding of the adolescent racers was postponed until the following May. By this time at an average age of 20 months, the snakes had made further growth.

Mr. Dwight R. Platt studied the changes in the male reproductive organs during the annual cycle at Harvey County. In racers recently emerged from hibernation he found the seminiferous tubules filled with Sertoli syncytium, but containing few germ cells. Spermatogonia proliferate in May and June. During the first half of July primary spermatocytes are the dominant cells in the seminiferous tubules. By early August spermatids are dominant and the first free spermatozoa are present. In late October spermiogenesis is essentially complete and the tubules are relatively empty before the snakes hibernate. During the season of activity the seminiferous tubules increase to approximately double their minimum diameter, reaching the maximum in August. Cyclic changes in size and secretory activity of the ductus deferens, ductus epididymis, and sexual segment of the renal tubules occur, with maximum size and secretory activity coinciding with the time of movement and storage of the spermatozoa. The latter are stored in both ductus deferens and ductus epididymis. Despite the short breeding season, a male racer has active sperm at all seasons.

Eggs

In accounts of the racer in the humid southeastern United States, Brimley (1903:261), Wright and Bishop (1915:160) and Tinkle (1959:195) mentioned the ease with which the eggs might be found and the superficial situations in which they were sometimes deposited. Both Wright and Bishop, and Tinkle made field studies in swamps, where presumably the subsoil was saturated with moisture and too wet for the eggs. Tinkle mentioned finding one clutch beneath a discarded newspaper and another beneath a small, thin board. Surface (1906:167) stated that in Pennsylvania the eggs were to be found in loose soil, in sawdust piles, or in decaying wood of hollow logs or trees. Clark (1949:249) stated that in northern Louisiana the eggs are laid in soft, moist soil such as may be found beside decaying logs. Minton (1944:457) found two clutches under flat stones on hillsides in Indiana. In the more arid climates of the far western states the species' habits are much different in this regard. Through many years of familiarity with C. c. mormon, I have never seen its eggs. Presumably nests in this part of the range are deep underground, most often in old burrows of the pocket gopher (Thomomys), which are so abundant that in many areas the soil is riddled with them. Burrows of the ground squirrels (Spermophilus sp.) and other small digging mammals also provide potential insulated nest sites with the favorably moderate temperatures and high humidities that the eggs of snakes require.