Natural History of Cottonmouth Moccasin, Agkistrodon piscovorus (Reptilia) Part 4

The "unidentified" category (Table 13) refers to jellylike ma.s.ses in the stomach or material in the intestine in which no scales, feathers, hair, or bones could be found. Most of the unidentifiable matter could be a.s.sumed to consist of remains of amphibians, since they leave no hard parts. If this a.s.sumption is correct, amphibians comprise about 40 per cent of the diet. Since intestinal contents were included, a volumetric a.n.a.lysis was not feasible. Therefore, the weight of each type of food item was estimated and the percentage by bulk calculated from it (Table 13).

Pieces of dead leaves and small sticks const.i.tuted most of the plant material found and presumably were ingested secondarily because they adhered to the moist skin of the prey, especially to fish and amphibians. However, some plant materials probably are eaten because they have acquired the odor of the prey. One cottonmouth contained a _Hyla cinerea_, several leaves, and five sticks from 37 to 95 millimeters long and from 12 to 14 millimeters in diameter.

Most reports in the literature state that gravid females do not feed, but four gravid females examined by me containing large, well-developed embryos also contained evidences of having recently fed. Two of them had scales of snakes in the stomach or intestine, one contained a six-inch _Lepomis_, and the other had hair in the intestine and the head and neck of an adult egret in the stomach.

MORTALITY FACTORS

Natural Enemies and Predators

Published records of other animals preying on cottonmouths or killing them are few. Reptiles more often than other cla.s.ses of vertebrates prey on the cottonmouth. McIlhenny (1935:44) reported on the scarcity of snakes in areas where alligators were present. Predation on cottonmouths by indigo snakes (_Drymarchon corais_) was reported by Conant (1958:153) and Lee (1964:32). Allen and Swindell (1948:6) obtained a photograph of a king-snake (_Lampropeltis getulus_) killing a cottonmouth but thought that moccasins are not eaten by _L. getulus_. However, one occasion reported herein shows that cottonmouths are eaten by king-snakes; and Clark (1949:252) reported finding 13 cottonmouths, along with other prey, in the stomach contents of 301 king-snakes (_L. g. holbrooki_) from northwestern Louisiana. Cannibalism is also common among cottonmouths. Klauber (1956:1058;1079) cited predation on cottonmouths by a blue heron (_Ardea herodias_) and a largemouth ba.s.s (_Micropterus salmoides_). Man is probably the greatest enemy of the cottonmouth.

Intentional killing, capturing, road kills, and alteration of the environment destroy large numbers.

Parasites and Diseases

Allen and Swindell (1948:12) listed several diseases and parasites of snakes and stated that "some moccasins captured in the woods are so poor and weak from parasitic infection that they can barely crawl." The only kind of ectoparasite found on captive cottonmouths in the course of my study was a snake mite, _Ophionyssus natricus_. An infestation of that mite was thought to be partly responsible for the death of one captive moccasin. Other moccasins spent increasing amounts of time in their water dish after they became infected with mites. Under natural conditions frequent swimming probably keeps cottonmouths nearly free of mites.

Endoparasites found included lung flukes, stomach nematodes, and tapeworms. Lung flukes (_Ochetosoma_ sp.) were found in 16 of 20 captive cottonmouths. Snails and frogs serve as intermediate hosts for various stages in the life cycle of these flukes. The high percentage of cottonmouths infested with flukes is indicative of the use of frogs as a major source of food. Less than ten flukes were usually observed in the snakes' mouths but occasionally more were seen. One snake was observed thras.h.i.+ng about in its cage for nearly an hour, after which time it died. Upon examination of the mouth, 32 flukes were found, most of which were located in the Jacobson's organs. Whether or not flukes caused the death is not known. Nematodes (_Kalicephalus_ sp.) were found in the stomach of each of several preserved specimens; most of these snakes had no food in their digestive tracts. In a high percentage of the moccasins, tapeworms (_Ophiotaenia_ sp.) were in the duodenum, in many instances so tightly packed as seemingly to prevent pa.s.sage of food. The importance of fish in the diet is reflected by the high percentage of snakes containing tapeworms. An unidentified cyst (?) about an inch long and containing two hooks on one end was found attached to the outer wall of the stomach of a cottonmouth. Yamaguti (1958) listed all the kinds of helminths known from cottonmouths.

Miscellaneous Causes of Death

Munro (1949:71-72) reported on the lethal effect of 10 per cent DDT powder on two young cottonmouths which were dusted with it to kill mites. Herald (1949:117) reported an equal effect caused by spraying a five per cent DDT solution in a room with several snakes. All but three large cottonmouths, which were under shelter at the time of spraying, were killed.

One individual that refused to eat was dissected soon after death, and a short piece of a branch on which two large thorns were located at 90 angles was found blocking the intestine at the posterior end of the stomach.

An unexpected and probably unusual circ.u.mstance caused the death of two captives. After cleaning a cage containing five cottonmouths and placing several mice in the cage for food, I noticed two of the snakes lying stretched out, partially on one side, and almost unable to move. At first I thought they had been bitten by other snakes which were in pursuit of the mice. The two died after two days. When a similar incident occurred in another cage, I removed the "bitten" snake and it fully recovered after 11 days. When the same symptoms were observed in a garter snake in another cage, I realized that in each instance the cage had been cleaned and fresh cedar chips placed in it immediately prior to observation of these symptoms. Fine cedar dust on the chips had evidently poisoned the snakes.

BEHAVIOR

Annual and Diel Cycles of Activity

In the days following emergence in spring, cottonmouths often endure uncomfortable and even dangerous temperatures in order to obtain food and mates. They are more sluggish at this time and more vulnerable to predation than later in the season when temperatures are optimal. Fitch (1956:463) found that copperheads in northeastern Kansas begin their annual cycle of activity in the latter part of April, when the daily maximum temperature is about 22 C. and the minimum is about 4 C., and become dormant in late October or early November, at which time the daily maximum temperature is about 15 and the minimum is about 0.

Indications are that in the northern part of its range the annual activity cycle of the cottonmouth resembles that of the copperhead in northeastern Kansas. Klimstra (1959:2) captured cottonmouths from April to October in southern Illinois. Barbour (1956:36) collected large numbers of them in early April in Kentucky and stated that they migrate from swamps to wooded hillsides in late August and early September.

Spring migrations begin after a few consecutive warm days in March. In northern Oklahoma cottonmouths have been found along the Verdigris River as early as March, suggesting that a few winter in crayfish holes and mammal burrows. The majority of individuals found in this area were at denning sites along cliffs above the river and emerged later than those near the river (Dundee and Burger, 1948:1-2). In Virginia cottonmouths have been seen as early as March 5 (Martin and Wood, 1955:237) and as late as December 4. They have been observed in migration from the swamps of the barrier beach to the mainland in late October and early November in southeastern Virginia (Wood, 1954a:159). According to Neill (1947:204), the cottonmouth tolerates lower temperatures than do most snakes in Georgia and is one of the last to go into hibernation. Allen and Swindell (1948:4) stated that cottonmouths usually bask during the mornings of the cooler months in Florida, but they mentioned nothing of denning such as occurs farther north. Although winter aggregations occur in the northern parts of the range, I have never seen such aggregations in the South. However, in one instance related to me by a reliable observer, seven cottonmouths were found together on a creek bank near the Gulf Coast in early spring.

During late summer and early autumn, fat is deposited in lobes in the lower abdomen in preparation for the period of winter quiescence. Gravid females usually do not feed so frequently or so much as other snakes, because they tend to become inactive as the ova develop. Whether or not females feed heavily after parturition and previous to denning is not known. Peaks of activity in autumn may be caused by final attempts to feed before denning and by the appearance of large numbers of newborn young. The young usually have from one to two months in which to feed before the advent of cold weather. According to Barbour's (_op.

cit._:38) findings, the young probably feed before hibernation because they grow substantially in winter. For those that do not feed, the rate of survival is perhaps much lower.

In preparation for winter, cottonmouths migrate inland, usually to dry forested hillsides where they den, commonly among rocks at the tops of bluffs, along with several other species of snakes. In such aggregations there is no hostility and each individual may derive benefit from contact with others by which favorable conditions of temperature and humidity are maintained.

Neill (1947:204) has found many specimens in winter by tearing bark from rotting pine stumps on hillsides overlooking lakes or streams. On cold days they evidently retreat below the surface, while on warm days they lie just below the bark or emerge and bask. Neill believes that the use of stumps by cottonmouths is an innate pattern of behavior, because of the large number of young-of-the-year found in such surroundings.

Cottonmouths were observed in winter also under logs and stumps by Allen (1932:17). I have twice observed cottonmouths crawling into crayfish burrows along the Gulf Coast of Texas, and suppose they are used as denning sites to some extent.

The diel cycle of activity of cottonmouths is of necessity closely related to the seasonal cycle. Since optimal temperatures determine activity, the diel cycle varies greatly from time to time. It has been well established that cottonmouths, like most other crotalids and many snakes of other families, prefer nocturnal to diurnal activity, even though the temperature may be less favorable at night. This preference is correlated with increased nocturnal activity of frogs and reptiles that const.i.tute the princ.i.p.al food supply.

During spring and autumn, activity is more restricted to the day and long periods of basking occur. However, as hot weather approaches, basking occurs mainly in the morning and evening and activity becomes primarily nocturnal. But, in well shaded, moist forests, cottonmouths feed actively in the daytime.

Availability of food also has an important influence upon activity.

Allen and Swindell (_op. cit._:5) stated that moccasins congregate around drying ponds and feed on dying fish until the moccasins can hold no more. They then usually stay nearby as long as food remains. In an area of the Stephen F. Austin Experimental Forest near Nacogdoches, Texas, many cottonmouths journey daily to and from a swamp and a dry field, evidently to feed on rodents inhabiting the area. Ten individuals captured along a snake-proof fence that was built 30 yards from the swamp were found lying coiled along the fence after 4:30 p.m., at which time the area was shaded. On another occasion, I captured a large cottonmouth that was feeding upon dying fish in a drying pool about 10:30 a.m. on August 19, 1962.

Because of the aquatic habits of the cottonmouth, relative humidity probably has little influence on the snake's activity. However, cottonmouths are more restricted to the vicinity of water in dry weather than during rains or muggy weather when many of their natural prey species also move about more freely. Increased activity on cloudy days may result from protection from long exposure to suns.h.i.+ne. Torrential rains and floods, such as those following hurricanes along the Gulf and Atlantic coasts of the southeastern United States, bring out quant.i.ties of snakes of all species. Rattlesnakes and cottonmouths in particular are killed by the thousands at these times because they seek shelter in human habitations. However, these are unusual circ.u.mstances and do not reflect voluntary activity as a result of preferences.

Thermal reactions of reptiles were cla.s.sified by Cowles and Bogert (1944) into several categories. For each species there is a basking and normal activity range limited by the voluntary minimum and voluntary maximum at which the animal seeks shelter. Beyond this normal range are the critical thermal minimum and critical thermal maximum (C. T. M.) at which effective locomotion is prevented. The lethal minimum and maximum are those temperatures at which short exposure produces irreparable damage, and death inevitably results. These cla.s.sifications are modified somewhat by seasonal or laboratory acclimation or by the physiological state of the animal. The C. T. M. of five cottonmouths was determined by placing each individual in an enclosed area and heating it with an infrared lamp. Cloacal temperatures were taken with a Schultheis quick-recording thermometer as soon as the snake could no longer right itself when placed on its back. All temperatures were in degrees Celcius. The C. T. M. averaged 39.2 (38.0 to 40.0). A temperature of 38.0 was lethal to one individual. These cottonmouths had been in captivity for nine months. The behavior of the snakes during heating resembled those instances described by Klauber (1956:382-387) for rattlesnakes. As the body temperature of the snakes rose past the optimum, each individual became disturbed and tried to escape from the enclosure. The snakes soon became frantic in their efforts to escape.

After about five minutes the mouth was opened and heavy, slow breathing was begun, accompanied by a loss of coordination and a slowing down of movements. The snakes writhed spasmodically for a few seconds and then lay still, usually with the mouth open. Recovery was begun by rolling on the belly and flicking the tongue, followed by movements of the head and then the body. Cottonmouths are rarely exposed to dangerously high temperatures owing to their semi-aquatic habits, but there are probably occasions when individuals reach the C. T. M. for the species.

Basking

Since activity, digestion, and gestation depend upon adequate internal temperatures, there must be a process by which these temperatures are attained and for an appropriate time maintained. Basking is important in this respect. The cottonmouths prefer to lie in a coiled position and, during basking, can usually be found beside bodies of water or on branches of dead trees overhanging the water. They are good climbers and have a prehensile tail that is frequently employed in descending from small branches. Since cottonmouths are semi-aquatic and are often exposed to temperatures that are lower than those of the air, they either must bask more often than terrestrial snakes or tolerate lower temperatures. Length of the period of basking is determined not only by amounts of insolation and temperature but also by the size of the snake.

A smaller snake can reach its optimum temperature more rapidly because of a higher surface-to-volume ratio. Another factor that may play a minor role in the rate of temperature change is the color of the snake.

The wide variation in color of cottonmouths probably affects rates of heat increase and loss due to direct radiation. Slight hormonal control of melanoph.o.r.es described in snakes by Neill and Allen (1955) also may exert some influence on the length of time spent basking. No rates of temperature increase or decrease are available for cottonmouths.

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 relations.h.i.+p 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 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.