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INTRODUCTION
INTRODUCTION
Objectives of the study here reported on were to: (1) learn as much as possible concerning the natural history and economic importance of the cottonmouth; (2) determine what factors limit its geographic distribution; (3) determine the role of the cottonmouth in its ecological community; and (4) compare the cottonmouth's life history with that of other crotalid snakes, especially the kinds that are most closely related to it. Twenty-five live cottonmouths were kept in the laboratory for the purpo
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ACKNOWLEDGMENTS
ACKNOWLEDGMENTS
For guidance in the course of my study, I am especially indebted to Professor Henry S. Fitch. For suggestions concerning the preparation of the manuscript, I thank Professor E. Raymond Hall. I am grateful to my wife, Janis, for her invaluable assistance and for typing the manuscript. For use of specimens in their care, I thank Professors William E. Duellman, University of Kansas; Robert L. Packard, formerly of Stephen F. Austin State College; W. Frank Blair, University of Texas; and William B. D
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SYSTEMATIC RELATIONSHIPS AND DISTRIBUTION
SYSTEMATIC RELATIONSHIPS AND DISTRIBUTION
Snakes of the genus Agkistrodon are relatively primitive members of the Crotalidae, which is one of the most specialized families of snakes. A majority of the pit-vipers are found in the Americas, but close relatives are found from extreme southeastern Europe through temperate Asia to Japan ( A. halys ) and southeastern Asia including Indonesia ( Agkistrodon and Trimeresurus ). Familial characters include: vertical pupil of the eye; facial pit present between the preoculars and loreal; scales us
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Color and Pattern
Color and Pattern
Color predominantly brown, ranging through pale reddish-brown or dark reddish-brown, brownish-green, to almost black; 10 to 17 irregular dark brown bands on paler brown ground color; young paler (some nearly salmon pink), retaining a vivid pattern throughout first year; pattern of most individuals nearly obliterated by third year; brilliance and dullness of predominant color correlated with molting cycle (skin especially bright and shiny immediately following shedding; tip of tail yellowish in j
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Scutellation
Scutellation
The scutellation of the cottonmouth closely resembles that of the other species of Agkistrodon . For example, the nine cephalic shields are characteristic of most species of Agkistrodon , as well as most other primitive crotalids and viperids, and most colubrids. Most individuals have an additional pair of large scales behind the parietals. The numbers of postoculars, supralabials, and infralabials are variable. On either side the postoculars (three in most specimens) are reduced to two in some
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Dentition
Dentition
Cottonmouths, like other pit-vipers, have their teeth reduced in number and have enlarged, highly specialized fangs. Small teeth occur on the palatine and the pterygoid in the upper jaw and on the dentary in the lower jaw. The dentary bone bears 17 curved teeth that decrease in size posteriorly. The palatine bears five small, strongly curved teeth, and the pterygoid bears 16 to 18 strongly curved teeth decreasing in size posteriorly. The numbers of teeth mentioned above in each instance refer to
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HABITAT AND LIMITING FACTORS
HABITAT AND LIMITING FACTORS
Although usually associated with swamps and lowlands along river bottoms, the cottonmouth lives in a variety of habitats ranging from salt marshes to cool, clear streams and from sea level to an altitude of 2300 feet. Shaded, moist areas either in or beside shallow waters are preferred, but cottonmouths occasionally wander as far as a mile from water. In the pine-oak forests of Nacogdoches County in eastern Texas cottonmouths and copperheads are probably the most abundant species of snakes. Spec
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Courtship and Mating
Courtship and Mating
A review of available literature indicates no records of courtship of the cottonmouth other than statements that breeding occurs in early spring. In a close relative, the copperhead (see Fitch, 1960:159-160), mating occurs almost any time in the season of activity but is mainly concentrated in the few weeks after spring emergence, at about the time when females are ovulating. Klauber (1956:692) concluded that along the southern border of the United States rattlesnakes normally mate in spring soo
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Reproductive Cycles
Reproductive Cycles
Many persons have assumed that gestation periods in snakes are the intervals between mating and parturition, and that mating and ovulation occur at approximately the same time. However, retention of spermatozoa and delayed fertilization indicate that copulation is not a stimulus for ovulation. A biennial reproductive cycle was found for the copperhead in Kansas (Fitch, 1960:162), the prairie rattler in Wyoming (Rahn, 1942:239) and in South Dakota (Klauber, 1956:688), the great basin rattler in U
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Embryonic Development
Embryonic Development
After ova are fertilized a three and one-half to four-month period of development begins which varies somewhat depending on the temperature. In almost every instance the ova in the right uterus outnumber those in the left. Embryos usually assume the serpentine form in the latter part of June and are coiled in a counterclockwise spiral with the head on the outside of the coil. At this time the head is relatively large and birdlike in appearance with conspicuous protruding eyes. Sex is easily note
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Birth of Young
Birth of Young
Accounts in the literature of 15 litters of cottonmouths fix the time of birth as August and September. Conant (1933:43) reported the birth of a litter in mid-July by a female that had been raised in captivity, and one female that I had kept in captivity for two months gave birth to a litter between October 19 and October 25. The conditions of captivity undoubtedly affected the time of birth in both instances. Wharton (1960:125-126) reported the birth and behavior of a brood of seven cottonmouth
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Number of Young per Litter
Number of Young per Litter
Records of from one to 16 young per litter have been reported (Ditmars, 1945:330; Clark, 1949:259), but the average is probably between six or seven. Most accounts in the literature present information on number of ova or embryos per female rather than the number of young. Size and age of the mother (Table 9) influence the number of ova produced. Allen and Swindell (1948:11) recorded three to 12 embryos in 31 cottonmouths varying in total length from 26 to 44 inches. An average of 6.5 embryos pe
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Population Composition
Population Composition
No investigator has yet analyzed the composition of a population of cottonmouths according to age, sex and snout-vent length. Barbour (1956:35) did sort 167 snakes into size classes, but did not determine sex ratio, size at sexual maturity, reproductive cycles, or snout-vent length. He recorded total lengths from which snout-vent lengths cannot be computed because of differential growth rates and different bodily proportions of the two sexes. I judge from my findings that he included immature in
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Reproductive Potential
Reproductive Potential
If data in Fig. 5 are representative of a natural population and if 61 per cent of the females are sexually mature, the reproductive potential can be estimated as follows: assuming a cohort of 1000 cottonmouths contains 530 females, 61 per cent of the females (323 individuals) probably are adults. If 42 per cent of these females produce 6.5 young per female in any season (Tables 8 and 9), 136 females will produce 884 young. But if 50 per cent of the adult females are reproductive (as would be as
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Size at Birth and Early Growth
Size at Birth and Early Growth
Size at birth depends on the health of the mother. According to Fitch (1960:182), many litters of copperheads born in captivity are stunted. Seven young cottonmouths (two males and five females) born in captivity were each 185 millimeters in snout-vent length and 40 millimeters in tail length. Weights of the three living young were 10.0, 10.1, and 11.1 grams. Another litter of five young measured by Richard S. Funk were larger, and differences in the proportions of the tail length and snout-vent
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The Umbilical Scar
The Umbilical Scar
The umbilical cord is broken at birth and the navel closes within a few days; but the scar, involving from two to four ventral scales, remains throughout life. Position of the scar was found by Edgren (1951:1) to be sexually dimorphic in the eastern hog-nose snake ( Heterodon platyrhinos ), but nothing has been published on this matter concerning the cottonmouth. Consequently, I counted the scales of several individuals from the anal plate, and there was no marked difference in the position of t
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Later Growth and Bodily Proportions
Later Growth and Bodily Proportions
The only records of growth increments in a natural population of cottonmouths are those in Table 11. The period of growth is mostly the period of activity, and differences are expected between northern and southern populations. As size increases, determination of growth rate becomes more difficult because age classes overlap in size. Growth of any individual depends not only on climate and food but also on disease and parasitism and the innate size potential. Stabler (1951:91) showed weight and
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The Shedding Operation
The Shedding Operation
Shedding of the skin is necessary to provide for growth and wear in snakes. The milkiness or bluing of the eyes, which causes partial blindness, marks the initial stage of shedding and is caused by a discharge of the exuvial glands that loosens the old stratum corneum from the layer below. In four to seven days the opaqueness disappears, and the snake sheds after an additional three to six days (Table 12). Young snakes first shed within a few days after birth and generally shed more frequently t
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Frequency of Shedding
Frequency of Shedding
Most of our knowledge concerning the frequency of shedding is based upon observations of captives. It is known that the intervals between exuviations are largely dependent upon the amount of food taken and the rate of growth. Unless laboratory conditions closely resemble those in the field, shedding frequencies in captives probably differ much from those of free-living snakes. Only two of my captives shed twice. The intervals between exuviations in the two snakes were eight and five months, last
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Methods of Obtaining Prey
Methods of Obtaining Prey
Food is obtained by a variety of methods depending on the type of food, age of the cottonmouth, and possibly other factors. Some captives lie in ambush and others crawl slowly in active search. At the first cue of possible prey, either by sight, scent, or differential temperature detection by the pit, the snake appears to become alert and flicks its tongue out at fairly rapid intervals. By means of the facial or loreal pit found in all crotalids, the snake is able to detect objects having temper
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Food and Food Preferences
Food and Food Preferences
The cottonmouth seems to be an opportunistic omni-carnivore, because it eats almost any type of flesh that is available, including carrion. It feeds primarily upon vertebrates found in or near water; but invertebrates and eggs have also been found in the diet. The only potential prey items that seem not to be normally eaten are bufonid toads and tadpoles. I have occasionally offered tadpoles and frogs to cottonmouths, but only the frogs were accepted. But, Stanley Roth kept a cottonmouth in capt
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Natural Enemies and Predators
Natural Enemies and Predators
Published records of other animals preying on cottonmouths or killing them are few. Reptiles more often than other classes 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
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Parasites and Diseases
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
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Miscellaneous Causes of Death
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
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Annual and Diel Cycles of Activity
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 do
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Basking
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
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Coiling
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, n
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Locomotion
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 substr
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Disposition
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 att
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Defense and Escape
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
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"Head Bobbing"
"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,
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Combat Dance
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
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Properties of 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. Kell
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Venom Yield and Toxicity
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 i
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Susceptibility of Snakes
Susceptibility of Snakes
Numerous experiments have been conducted to determine the susceptibility of various snakes to venom. The majority of these experiments were performed to learn whether or not venomous snakes were immune to their own poison. Conant (1934:382) reported on a 30-inch cottonmouth that killed two Pacific rattlesnakes and another cottonmouth. One rattlesnake was bitten on the tail and the other on or near the head and partially swallowed. Gloyd (1933:13-14) recorded fatal effects to a rattlesnake from t
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Effects of the Bite
Effects of the Bite
Factors determining the outcome of snakebite are: size, health, and species of snake; individual variation of venom toxicity of the species; age and size of the victim; allergic or immune responses; location of the bite; and the amount of venom injected and the depth to which it is injected. The last factor is one of the most variable, owing to (1) character and thickness of clothing between the snake and the victim's skin, (2) accuracy of the snake's strike, and (3) size of the snake, since a l
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Treatment
Treatment
Perhaps one of the most important factors in the outcome of snakebite is the treatment. Because of the variable reactions to snakebite, treatment should vary accordingly. Many methods have been proposed for treating snakebite, and there is disagreement as to which is the best. The list of remedies that have been used in cases of snakebite includes many that add additional injury or that possibly increase the action of the venom. The use of poultices made by splitting open living chickens and the
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Case History of a Bite
Case History of a Bite
On July 29, 1963, at 8:20 a.m., I was treating a nine-month-old cottonmouth for mites. As I dropped the snake into a sink, it twisted its head and bit the tip of my right middle finger with one fang. The fang entered just under the fingernail and was directed downward, the venom being injected about five millimeters below the site of fang penetration. After placing the snake back in its cage, I squeezed the finger once to promote bleeding, wrapped a string around the base of the finger, and drov
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Snakebite in the United States
Snakebite in the United States
Many estimates have been made of the number of bites of poisonous snakes that occur annually in the United States. The occurrence of poisonous snakebite has been nearly as badly underestimated as fatal results of their envenomations have been overrated. For important data on number of persons bitten by poisonous snakes in the United States, see the following: Allen and Swindell (1948:15); Githens (1935:172); Klauber (1956:811); Parrish (1963); Sowder and Gehres (1963:973); Stimson and Engelhardt
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SUMMARY
SUMMARY
In my study, 306 living and preserved cottonmouths were examined. This species occurs throughout the coastal plains of the southeastern United States, usually at altitudes of less than 500 feet but occasionally up to altitudes of more than 2000 feet. Two subspecies are recognized: the eastern cottonmouth, A. p. piscivorus , occurring from extreme eastern Mississippi to southeastern Virginia and Florida; and the western cottonmouth, A. p. leucostoma , occurring from eastern Mississippi northward
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LITERATURE CITED
LITERATURE CITED
Allen, E. R. 1937. Florida snake venom experiments. Florida Acad. Sci., 2:7 pp. Allen, E. R. , and Swindell, D. 1948. Cottonmouth moccasin of Florida. Herpetologica, 4:1-16 (first supplement). Allen, M. J. 1932. A survey of the amphibians and reptiles of Harrison County, Mississippi. Amer. Mus. Novit., 542:1-20. Anderson, P. 1941. The cottonmouth in northern Missouri. Copeia, 1941(3):178. 1945. New herpetological records for Missouri. Bull. Chicago Acad. Sci., 7(5):271-275. Anon. 1953. Snake col
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UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY
UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY
Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. When copies are requested from the Museum, 25 cents should be included (for each 100 pages or part thereof) for the purpose of defraying the costs of
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