Part 8

Coiling

While inactive the cottonmouth spends most of its time lying in a coiled position with the tail outermost, with the body usually wound into about one and one-half cycles, and the head and neck in a reversed direction forming a U- or S-shaped loop. From this position the snake is able to make a short strike or a hasty getaway if necessary. In my opinion this position is used primarily for basking or resting and only secondarily for feeding. Most individuals appear to pursue their prey actively, not lying in ambush for the approaching prey to the extent that most other crotalids do.

Many of the cottonmouths that I kept in captivity were observed in a coiled position for periods up to three or four days. Under natural conditions, however, they are more active. Young cottonmouths are inclined to remain in a coiled position for longer periods than older individuals.

Locomotion

Four distinct types of locomotion have been described in snakes: horizontal undulatory, rectilinear, sidewinding, and concertina (Klauber, 1956: 331-350). Most snakes are capable of employing two or more of these types of progression, at least to a certain degree; but horizontal undulatory locomotion is the most common method used by the majority of snakes, including the cottonmouth. In this method the snake's body is thrown into lateral undulations that conform with irregularities in the substrate. Pressure is exerted on the outside and posterior surface of each curve, thus forcing the body forward.

Rectilinear locomotion is more useful to large, thick-bodied snakes which use this method of progression, chiefly when they are prowling and unhurried. This method depends upon the movement of alternate sections of the venter forward and drawing the body over the ventral scales resting on the substratum by means of muscular action. This mode of locomotion was most frequently observed in captive cottonmouths when they were crawling along the edge of their cages, especially when they were first introduced to the cages and toward the end of the shedding process. The other two types of locomotion, sidewinding and concertina, have not, to my knowledge, been observed in the cottonmouth.

Both the cottonmouth and the cantil have definite affinities for water and are as likely to be found in water as out of it. Copperheads and rattlesnakes, although not aquatic, are good swimmers. When swimming, a motion resembling horizontal undulatory progression is used.

Disposition

The number of different opinions expressed in the literature concerning the cottonmouth's disposition is not at all surprising. As with any species there is a wide range of individual temperament, which is affected by many factors. The cottonmouth is considered by some writers to be docile while others consider it to be highly dangerous. Allen and Swindell (1948:7) described the variability in temperament, even among individuals. They wrote: "On rare occasions, moccasins are found which will attack. A perfectly docile snake will turn and bite viciously without any apparent reason." They also recounted a case in which a cottonmouth was kept as a pet for six years, being allowed the freedom of the house. Smith and List (1955:123) found them "... surprisingly docile in the gulf region [Mississippi], displaying none of the pugnacity of more northern cottonmouths." Smith (1956:310) stated: "Unlike the copperhead, cottonmouths are pugnacious; their powerful jaws, long fangs, vicious disposition and potent venom make them a very dangerous animal."

My own observations are in general agreement with the statements of Allen and Swindell (loc. cit.). In my encounters with cottonmouths, I have never found any aggressive individuals except for three juveniles that were born in captivity. In their first three days in the laboratory these juveniles were observed to strike repeatedly whenever anyone entered the room. After this short period of aggressiveness, however, they slowly became more docile. The disposition shown by the newborn young is clearly an innate behavioral pattern that undoubtedly has a direct relationship to survival. The majority of cottonmouths that I have approached in the field have moved swiftly to seek refuge in nearby water; a few have remained motionless as I approached, and one showed the typical threat display. Upon capture and handling, they react similarly to other pit-vipers by opening and closing the mouth and erecting the fangs in an attempt to bite. They often bite through the lower jaw and eject venom at this time as well as when the mouth is open. Of more than a dozen individuals kept in captivity, four were particularly difficult to handle whereas another was extremely docile. It was almost never found in aggregations with the other snakes and did not struggle or attempt to bite when handled. The majority remained unpredictable in disposition, usually appearing docile and lazy but capable of extremely rapid movements when disturbed.

Defense and Escape

The typical threatening posture of rattlesnakes is all but lacking in the cottonmouth, which relies primarily on concealing coloration or nearness to water for escape. When approached, it usually plunges into nearby water or remains motionless with the head held up at a 45 deg. angle and the mouth opened widely exposing the white interior. The tail is sometimes vibrated rapidly and musk is expelled. This threat display is unique to cottonmouths; although it does not attract as much attention as the display of rattlesnakes, it is probably an effective warning to most intruders at close range.

Neill (1947:205) reported one case in which a cottonmouth used the "body blow" defense, described for Crotalus by Cowles (1938:13), when approached by a king-snake, Lampropeltis getulus. In this unusual posture the anterior and posterior portions of the body are held against the ground and the middle one-fourth to one-third of the body is lifted up and used in striking the intruder. This same defense posture also was observed in rattlesnakes when presented with the odor of the spotted skunk, Spilogale phenax. However, the "king-snake defense posture" is probably not a well-established behavioral pattern in the cottonmouth, for it sometimes feeds upon king-snakes. I observed the killing and devouring of a cottonmouth by a speckled king-snake, L. g. holbrooki; the only attempts to escape were by rapid crawling and biting.

Cottonmouths often squirt musk as a defensive action. The tail is switched back and forth, and musk is emitted from glands on each side of the base of the tail. The fine jets of musk are sprayed upward at about 45 deg. angles for a distance of nearly five feet. How often this defense mechanism is used against other animals is not known, but the musky odor can frequently be detected in areas where cottonmouths are common. The odor is repulsive and, if concentrated, can cause nausea in some individuals. To me, the scent is indistinguishable from that of the copperhead.

"Head Bobbing"

"Head bobbing" in snakes has been described frequently in the literature, and many interpretations have been advanced to explain its occurrence. One of the earlier accounts was that of Corrington (1929:72) describing behavior of the corn snake, Elaphe guttata. Characteristic bobbing occurred when the snake was cornered, and seemingly the purpose was to warn or frighten foes. Neill (1949:114-115) mentioned the jerking or bobbing of the head in several species of snakes including the cottonmouth, and remarked that "it is apparently connected with courtship and with the recognition of individuals." According to Munro (1950:88), "head bobbing" appears to be a sign of annoyance in some instances but is usually concerned with reproduction and individual recognition. Richmond (1952:38) thought that many types of head movements among not only reptiles but also birds and some mammals are a result of poor vision and serve "to delimit and orient an object that for lack of motion is otherwise invisible." Head movements undoubtedly occur in animals to facilitate accommodation, but it is obvious from Richmond's conclusions that he has never observed "head bobbing" in snakes. The term itself is grossly misleading and should be discarded. Mansueti (1946:98) correctly described the movements as spastic contractions of the body. I have observed numerous instances of these movements in cottonmouths, copperheads, and rat snakes (Elaphe obsoleta); and in no case has the movement resembled a head bob as is described in lizards and other animals. The movement appears to be a result of a nervous or sexually excited state and consists of highly spastic contractions confined to the anterior part of the snake most of the time but affecting the entire body on some occasions.

I found the response to be most common among cottonmouths in confinement when food was introduced to a cage containing several individuals (increasing the tendency to strike at a moving object) and when an individual was placed back in the cage after being handled. At these times the snakes that were inactive began to jerk for a few seconds. When the snake is in this seemingly nervous state, the same response is elicited by another snake crawling over it. At other times the movement of one individual causes no such response. The jerking movements appear to be released by the recognition of a nervous state in another individual and may serve to protect the jerking individual from aggressive advances of the former.

Where courtship is involved, the jerking motions are made in conjunction with writhing of the male and do not result from the same type of releaser described above.

Combat Dance

The so-called combat dance between male snakes has long been known, but its significance is still poorly understood. It was for many years believed to be courtship behavior until the participants were examined and found to be males. Carr and Carr (1942:1-6) described one such instance in two cottonmouths as courtship. In their observations, as well as those of others, copulation was never observed following the "dance" but was assumed to be the ultimate goal. After the discovery that only males participated, it was suggested that combat involved competition for mates, but the "dance" has been observed at times other than the breeding season (Ramsey, 1948:228).

Shaw (1948:137-145) discussed the combat of crotalids in some detail but drew no conclusions as to the cause of the behavior. Lowe (1948:134) concluded with little actual evidence that combat among male snakes is solely for territorial purposes. Shaw (1951:167) stated that combat may occur as a possible defense against homosexuality. One case of homosexual mating among cottonmouths was reported (Lederer, 1931:651-653), but the incomplete description seems to be of normal courtship procedure except that the "female" tried to avoid the male. Two instances of combat observed between timber rattlesnakes (C. h. horridus) by Sutherland (1958:23-24) were definitely initiated because of competition for food. More observations are needed before the significance of the combat can be fully understood.

THE VENOM

Properties of the Venom

The venom and poison apparatus have developed primarily as a means of causing rapid death in small animals that are the usual prey. As a protective device against larger enemies, including man, the venom may have some value; but this was probably unimportant in the evolution of the poison mechanism. A secondary function of the venom is to begin digestion of tissues of the prey. Since food is swallowed whole, injection of digestive enzymes into the body cavity enhances digestion of the prey.

Kellogg (1925:5) described venom as a somewhat viscid fluid of a yellowish color and composed of 50 to 70 per cent proteins, the chief remaining components being water and carbohydrates, with occasional admixtures of abraded epithelial cells or saprophytic microorganisms. Salts, such as chlorides, phosphates of calcium, magnesium, and ammonium, occur in small quantities. Each of the components of snake venom has a different effect on the body of the victim. It was at first believed that there were two types of venoms: neurotoxic, which acts upon nervous tissue; and haemotoxic, which acts on blood and other tissues. It has since been found that venoms are composed of varying mixtures of both types. Fairley (1929:301) described the constituents of venom as: (1) neurotoxic elements that act on the bulbar and spinal ganglion cells of the central nervous system; (2) hemorrhagins that destroy the lining of the walls of blood vessels; (3) thrombose, producing clots within blood vessels; (4) hemolysins, destroying red blood corpuscles; (5) cytolysins that act on leucocytes and on cells of other tissues; (6) elements that retard coagulation of the blood; (7) antibactericidal substances; and (8) ferments that prepare food for pancreatic digestion. Elapid snakes tend to have more of elements 1, 4, and 6 in their venoms, while viperids and crotalids, of which the cottonmouth is one, have higher quantities of elements 2, 3, and 5. Kellogg (loc. cit.) stated that venom of cottonmouths contains more neurotoxin than that of rattlesnakes and not only breaks down the nuclei of ganglion cells but also produces granular disintegration of the myelin sheath and fragmentation of the conducting portions of nerve fibers.

Thus, venoms contain both toxic elements and non-toxic substances that promote rapid spreading of the venom through the body of the victim. Jacques (1956:291) attributed this rapid spreading to the hyaluronidase content of venoms.

Venom Yield and Toxicity

One of the most important yet undeterminable factors of the gravity of snakebite is the amount of venom injected into the victim. Since this volume varies considerably in every bite, attempts have been made to determine the amount and toxicity of venom produced by each species of poisonous snake. Individual yield is so variable that a large number of snakes must be milked in order to determine the average yield. Even then there remains an uncertainty as to how this amount may compare with that injected by a biting snake.