Natural History of Cottonmouth Moccasin, Agkistrodon piscovorus (Reptilia)
Part 6
It can be seen from the above examples that toxicity of venoms and the resistance of the animal receiving an injection of venom is highly variable. Possibly the venom of each species of snake has greatest effect on animals of the particular group relied on for food by the snake. If that is so, the venom of cottonmouths would be expected to be more toxic when tested on fish, reptiles, and amphibians than on birds and mammals. Likewise, the venom of most species of rattlesnakes would be expected to be more virulent when injected into mammals than when injected into lower vertebrates. But, according to Netting (1929:108), species of rattlesnakes that prey on cold-blooded animals, which are less susceptible to venoms than warm-blooded animals, are thought to have highly toxic venoms. This explanation accounts for the powerful venom of _Sistrurus catenatus_; and, in this respect, venom of cottonmouths should be highly toxic also. However, no clear-cut trends have been shown in most cases. Allen (1937) injected 250-gram guinea pigs with 4 milligrams of venom of various poisonous snakes. Survival time was recorded in order to indicate the relative potency of the venoms. Of 16 such tests _C. adamanteus_ held places 1, 2, 3, 12, and 16; _Bothrops atrox_ held places 4, 9, 10, and 13; and _A. piscivorus_ held places 5, 7, 8, and 15. Places 6, 11, and 14 were held by three individuals of different species. No relationship to size or sex was indicated by the results of this experiment.
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 the bite of a cottonmouth. He also reported on the observations of three other crotalids bitten by themselves or other snakes, from which no harmful effects were observed. Allen (1937) injected several snakes with dried cottonmouth venom which was diluted with distilled water just before each injection. Four cottonmouths receiving 9, 18, 19, and 20 milligrams of venom per ounce of body weight survived, while another receiving 18.7 milligrams per ounce died after three hours. A specimen of _S. miliarius_ receiving 8.3 milligrams per ounce died in about ten hours, while a _C. durissus_ receiving 12.5 milligrams per ounce succumbed in 45 minutes. An alligator receiving 6 milligrams per ounce died in 14 hours. Even the snakes that survived showed some degree of swelling.
The studies of Keegan and Andrews (1942:252) show that king-snakes are sometimes killed by poisonous snakes. A _Lampropeltis calligaster_ injected with _A. contortrix_ venom (0.767 milligrams per gram) died five days following the injection. This amount was more than twice the amount of _A. piscivorus_ venom injected into a _L. getulus_ by Allen (1937) in which the snake showed no ill effects. Keegan and Andrews (_loc. cit._) stated that success in overpowering and eating poisonous snakes by _Lampropeltis_ and _Drymarchon_ may be due to the ability to avoid bites rather than to immunity to the venom. However, Rosenfeld and Glass (1940) demonstrated that the plasma of _L. g. getulus_ had an inhibiting effect on the hemorrhagic action on mice of the venoms of several vipers.
One of the more extensive studies on effects of venoms on snakes is that by Swanson (1946:242-249). In his studies freshly extracted liquid venom was used. His studies indicated that snakes are not immune to venom of their own kind or to closely related species. Copperhead venom killed copperheads faster than did other venoms but took more time to kill massasaugas, cottonmouths, and timber rattlers. However, most of the snakes were able to survive normal or average doses of venom although they are not necessarily immune to it.
One of the major problems in comparing the data on toxicity of venom in studies of this type is that no standard method of estimating toxicity has been used. Swanson's (_loc. cit._) amount of venom equalling one minim (M.L.D.?) ranged from 0.058 to 0.065 cubic centimeters. There were no different values given for each species, but the time that elapsed from injection of the venom to death represented the toxicity. There also was no attempt in his study to convert the amount of venom used into a ratio of the volume of venom per weight of snake, making the results somewhat difficult to interpret. Additional work in this field should provide for many injections into many individuals of several size classes. The studies to date have been on far too few individuals to allow statistical analyses to be accurate.
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 large snake can deliver more venom and at a greater depth than can a small snake.
Pope and Perkins (1944) demonstrated that pit-vipers of the United States bite as effectively as most innocuous snakes and that a careful study of the bite may reveal the location of the pocket of venom, size of the snake, and possibly its generic identity (see Dentition). The bite pattern of the cottonmouth as well as the other crotalids showed the typical fang punctures plus punctures of teeth on both the pterygoid and mandible. Even so, a varying picture may be presented because from one to four fang marks may be present. At times in the fang-shedding cycle three and even four fangs can be in operation simultaneously.
Various authors have attributed death of the prey to the following causes: paralysis of the central nervous system, paralysis of the respiratory center, asphyxiation from clotting of the blood, stoppage of the heart, urine suppression due to crystallized hemoglobin in the kidney tubules, dehydration of the body following edema in the area of the bite, or tissue damage. Mouths of snakes are reservoirs for infectious bacteria, which are especially prolific in damaged tissue. Bacterial growth is aided by the venom which blocks the bactericidal power of the blood.
Three grades in the severity of snakebite (I, minimal; II, moderate; and III, severe) were described by Wood, Hoback, and Green (1955). Parrish (1959:396) added a zero classification to describe the bite of a poisonous snake in which no envenomation occurred. Grade IV (very severe) was added by McCollough and Gennaro (1963:961) to account for many bites of the eastern and western diamondback rattlesnakes.
The first symptom of poisonous snakebite is an immediate burning sensation at the site of the bite. Within a few minutes the loss of blood into the tissues causes discoloration. Swelling proceeds rapidly and can become so great as to rupture the skin. Pain is soon felt in the lymph ducts and glands. Weakness, nausea, and vomiting may ensue at a relatively early stage. Loss of blood into tissues may spread to the internal organs. In conjunction with a rapid pulse, the blood pressure and body temperature can drop. Some difficulty in breathing can occur, especially if large amounts of neurotoxin are present in the venom. In severe cases the tension due to edema obstructs venous and even arterial flow, in which case bacteria may multiply rapidly in the necrotic tissue and gangrene can occur. Blindness due to retinal hemorrhages may occur. Symptoms of shock may be present after any bite.
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 use of alcohol, potassium permanganate, strychnine, caffeine, or injection of ammonia have no known therapeutic value, and may cause serious complications. The most important steps in the treatment of snakebite are to prevent the spread of lethal doses of venom, to remove as much venom as possible, and to neutralize the venom as quickly as possible.
It is generally agreed that the first step in snakebite treatment should be to place a ligature above the bite to restrict the flow of venom, and also to immobilize the patient as much as possible. The ligature should be loosened at least every fifteen minutes. The next steps are sterilization of the skin and the making of an incision through the fang punctures. As pointed out by Stahnke (1954:8), the incision should be made in line with the snake's body at the time of the bite, so as to account for the rearward curvature of the fangs and possibly to reach the deposition of venom. Many instruction booklets and first-aid guides have specified the length and depth of incision to be made, but the actual size and depth of the cut should depend upon the location of the bite. An "X" cut or connection of the fang punctures is likely to facilitate the spread of the venom. No cut should be made that would sever a large blood vessel or ligament.
Extensive damage is often caused by well-meaning individuals whose attempts at first aid result in brutally deep incisions and tourniquets applied too tightly and for too long a period of time; the resultant damage in many instances exceeds that of the bite itself (Stimson and Engelhardt, 1960:165). Stimson and Engelhardt also think that time should be sacrificed to surgical cleanliness, and incisions should not be made if a hospital can be reached within an hour.
The ligature-cryotherapy (L-C) method proposed by Stahnke (1953) has been severely criticized by other workers. He stated that the ligature should be tight enough to restrict completely the flow of venom until the temperature of the area can be lowered sufficiently to prevent any action of the venom. After 10 minutes the ligature may be removed and the bitten area kept immersed in a vessel of crushed ice and water. If the envenomized member is to be treated for more than four hours (which is the case with almost all pit-viper bites), it should be protected by placing it in a plastic bag. The venom action should be tested after 12 or more hours. This consists of a brief warming period to determine whether or not the action of the venom can be felt. The patient should be kept warm at all times; and the warming at the termination of treatment should be done gradually, preferably by allowing the water to warm slowly to room temperature.
Advocates of the L-C method warn against making incisions unless they are absolutely necessary, the theory being that each cut permits additional bacterial infection and does little good in removing venom. However, McCollough and Gennaro (1963:963) demonstrated that, in bites where the fangs had only slightly penetrated the skin, more than 50 per cent of the venom was removed in some instances if suction was started within three minutes after the injection. With deeper injection the amount of venom recovered sometimes reached 20 per cent of the dose. Stahnke suggested that an incision be made at the site of the bite only after the site has been refrigerated for at least 30 minutes.
Stimson and Engelhardt (_loc. cit._) stated that two constricting bands should be used between the bite and the body and that cracked ice in a cloth should be applied to the bite before reaching a hospital. In addition, they suggested the following procedure. Rings of incisions should follow the swelling, and suction should continue for several hours. After the edema has receded, the limb should be wrapped in a towel containing crushed ice. Antivenin should be given only in severe cases. Calcium gluconate and gas gangrene antitoxin as well as antibiotics are helpful.
The most recent and up-to-date summary of snakebite treatment is that by McCollough and Gennaro (1963). Following is a brief summary of their suggestions:
1. Immobilization--Systemic immobilization is effected by body rest and locally by splinting the bitten area.
2. Tourniquet--A lightly occlusive tourniquet during a 30- to 60-minute period of incision and suction would seem to possess some advantages. In severe cases where medical attention is hours away, a completely occlusive tourniquet may be necessary to prevent death. Sacrifice of the extremity may be necessary for the preservation of life.
3. Incision and suction--Suction should begin three to five minutes after injection of venom if symptoms of poisoning are present. Incisions one-fourth inch to an inch long across each fang mark should be made in order to open the wound for more efficient suction. Multiple incisions are not useful for the removal of venom but may be employed under hospital conditions to reduce subcutaneous tensions and ischemia.
4. Cryotherapy--An ice cap over the site of the bite for relief of pain would seem to be permissible, especially prior to the administration of antivenin. It must be remembered that cooling during the administration of the antivenin radically reduces the access of the antiserum to the bite area.
5. Antivenin--Antiserum is the keystone to the therapy of snakebite. Careful evaluation of the severity of the bite and the patient's sensitivity should be made before the use of antivenin. In Grade II (moderate) bites, the intramuscular injection on the side of the bite may suffice. In Grades III (severe) and IV (very severe), shock and systemic effects require intravenous injection. In bites producing symptoms of this severity, antivenin must be given in amounts large enough to produce clinical improvement. Ten to 20 units may be necessary to prevent the relapse that sometimes occurs after small doses of antivenin. Permanent remission of swelling and interruption of necrosis are the therapeutic end point in the clinical use of the antiserum.
In all cases of snakebite where there is any doubt as to the snake's identity, it should be killed if possible and taken to the hospital for positive identification. In many instances of actual bites by poisonous snakes the only treatment needed was an injection of tetanus antitoxin or toxoid and sedation, because physical examination revealed no indication of poisoning (Stimson and Engelhardt, _loc. cit._).
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 drove to Watkins Memorial Hospital on the University of Kansas campus. I began to feel a burning sensation in the tip of the finger almost immediately. Upon my arrival at the hospital, an additional ligature was placed around my wrist. At 8:30 a.m. a small incision was made in the end of the finger, which by this time was beginning to darken at the point of venom deposition. I sucked on the finger until 8:35 a.m., when a pan of ice water that I had requested was brought to me. No pain was felt except that caused by the ice. Fresh ice was added as needed to keep the temperature low. By 9:30 a.m. the finger had swollen and stiffened. At 10:00 a.m. the swelling had progressed to the index finger and back of the hand. I experienced difficulty in opening and closing the hand. Blood oozed slowly from the incision. A dull ache persisted and about every two to four minutes a sharp throb could be felt until nearly 11:00 a.m., when the pain diminished. The rate and intensity of throbbing increased whenever the hand was removed from the ice bath for more than a few seconds. Although only the hand was immersed, the entire forearm was cold. Pain was felt along the lymphatics on top of the arm when it was touched, and by 1:00 p.m. a slight pain could be felt in the armpit. Since swelling and pain were almost nonexistent by 2:00 p.m., I was permitted to leave. After walking to a nearby building, I again felt a burning sensation as the hand warmed. I made another ice bath and again immersed the hand in it until 4:10 p.m., at which time it was removed from the water. The pain and swelling began anew, and the hand was placed back in an ice bath from 5:30 p.m. until about 7:30 p.m. At this time cryotherapy was discontinued. From 10:00 p.m. to 12:00 midnight my legs twitched periodically, and pain could be felt in both armpits. A slight difficulty in breathing also was experienced for a short time. The acute pain and burning sensation remained in the finger until the following morning, but swelling progressed only as far as the wrist. The only discomfort that day was in the finger. The tip was darkened, the entire first digit red and feverish, and the lymphatics still painful when touched. By the third day the swelling had regressed. The incision itself was the main cause of discomfort, and the soreness at the site of the bite persisted for at least four days.
Although the L-C method of snakebite treatment has been vigorously attacked by many, there is still need of much more data before it can be unequivocally condemned or praised. It was preferred in the treatment of this bite because: I knew that envenomation was minimal and that there would be no need for antivenin; only one fang of a snake less than one foot long had entered the tip of the finger; the snake had bitten three frogs in the previous two days and had possibly used up a considerable amount of its venom; the venom was deposited at such a shallow depth that at least a portion of it could be removed by suction; and the wound bled freely even before suction was applied. The ice water was uncomfortably cold but was not cold enough to cause frostbite, a major objection to the L-C method. Ideally, fresh ice should be added little by little to replace that which is melting, and the immersed area should be protected from the water by a plastic bag. Pain and swelling can be minimized by cryotherapy, but I would recommend its use only in cases of mild poisoning such as the one described herein.
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 (1960:153); Swaroop and Grab (1956:441); Swartzwelder (1950:579); Willson (1908:530); and Wood (1954b:937).
Judging from estimates made in several states, the number of poisonous snakebites in the United States would be about 5000 per year. In the region where the cottonmouth occurs there are approximately 2000 persons bitten annually by poisonous snakes. Of these approximately 39 per cent are copperhead bites, 30 per cent each are cottonmouth and rattlesnake bites, and I per cent are coral snake bites. These percentages vary considerably from place to place, because of the distribution and abundance of the eight species of poisonous snakes whose ranges overlap that of the cottonmouth.
According to Parrish (1963), about 14 people die of snakebite each year in the United States. Of these deaths, about 6.6 per cent are attributable to cottonmouths, 77.0 per cent to rattlesnakes, and 1.6 per cent to coral snakes; 14.8 per cent are unidentified. Almost half of the fatalities are in persons less than 20 years of age, the high mortality rate being partially due to the greater ratio of venom to body weight.
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 to southern Illinois and Missouri and westward to central Texas. Intergradation occurs in eastern Mississippi.
The northern edge of the range is probably limited by low temperatures in winter, and the western edge by lack of available habitat resulting from insufficient precipitation. Old records of occurrence indicate that the range has decreased in the last 100 years. The species inhabits mostly areas where water is found, but at times wanders a mile or more from the nearest water.
The ground color is predominantly a brown, but varies from a brownish-green to almost black with a pattern of 10 to 17 irregular bands of a darker shade of brown. The pattern is better defined in the eastern subspecies than in the western.
The scutellation resembles that of other species of _Agkistrodon_. In the specimens examined supralabials ranged from 7 to 9, and infralabials from 8 to 12. The number of dorsal scale rows on the neck, at mid-body, and immediately anterior to the anus is relatively constant at 27-25-21, respectively. Ventral scales of 34 males averaged 134.4 (128 to 139), and those of 48 females 133.5 (128 to 137). The number of caudal scales showed some degree of sexual dimorphism; the average was 45.4 (41 to 50) in 34 males and 42.6 (39 to 49) in 44 females. In general, caudal scales on the basal half of the tail are undivided, whereas those on the distal half are divided. No marked geographical variation was found in any scale character.
The poison fangs vary in length from 1.3 per cent of snout-vent length in juveniles to 1.0 per cent in large adults. Fangs of captive cottonmouths were shed and replaced at intervals of about 21 days, but the interval was variable. Relationships in distance between the base of fangs and between fang punctures in an actual bite indicate that examination of the wound does not provide a good basis for judging accurately the size of the snake that inflicted the bite.