Rattlesnake

From Wikipedia, the free encyclopedia
Jump to: navigation, search
Rattlesnake
Crotalus cerastes
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Class: Reptilia
Order: Squamata
Suborder: Serpentes
Family: Viperidae
Subfamily: Crotalinae
Genera

Crotalus Linnaeus, 1758
Sistrurus Garman, 1883

Rattlesnakes are a group of venomous snakes of the genera Crotalus and Sistrurus[1] of the subfamily Crotalinae ("pit vipers"). There are 32 known species of rattlesnakes, with between 65-70 subspecies,[2] all native to the Americas, ranging from southern Alberta and southern British Columbia in Canada to Central Argentina.

Rattlesnakes are predators who live in a wide array of habitats, hunting small animals such as birds and rodents. They kill their prey with a venomous bite, rather than by constricting. All rattlesnakes possess a set of fangs with which they inject large quantities of hemotoxic venom. The venom travels through the bloodstream, destroying tissue and causing swelling, internal bleeding, and intense pain. Some species, such as the Tiger Rattlesnake and the Mojave Rattlesnake, additionally possess a neurotoxic component in their venom that causes paralysis and other nervous symptoms.

The threat of envenomation, advertised by the loud shaking of the titular noisemaker ("rattle") at the end of their tail, deters many predators. However, rattlesnakes fall prey to hawks, weasels, king snakes, and a variety of other species. Rattlesnakes are heavily preyed upon as neonates, while they are still weak and mentally immature. Very large numbers of rattlesnakes are killed by humans. Rattlesnake populations in many areas are severely threatened by habitat destruction, poaching, and extermination campaigns.

Rattlesnake bites are the leading cause of snakebite injuries in North America. However, rattlesnakes rarely bite unless provoked or threatened; and if treated promptly, the bites are rarely fatal.

Names[edit]

Rattlesnakes receive their name from the rattle located at the end of their tails, which, when shaken, makes a loud rattling noise that deters predators or serves as a warning to passersby.[3] The scientific name Crotalus derives from the Greek, κρόταλον, meaning "castanet".[4] The name Sistrurus is the Latinized form of the Greek word for "tail rattler" (Σείστρουρος, Seistrouros) and shares its root with the ancient Egyptian musical instrument, the sistrum, a type of rattle.[5]

Ecology[edit]

Range and habitat[edit]

Western diamondback rattlesnake (Crotalus atrox), responsible for the majority of venomous snakebites in North America, coiled in defensive posture with rattle erect

Rattlesnakes are native to the Americas, living in diverse habitats from Southwestern Canada to Central Argentina. The large majority of species live in the American Southwest and Mexico. Four species may be found east of the Mississippi river, and two in South America. In the United States, the states with the most types of rattlesnakes are Texas and Arizona.

Rattlesnakes are found in almost every type of habitat that is capable of supporting terrestrial ectothermic vertebrates; but individual species can have extremely specific habitat requirements, only able to live within certain plant associations in a narrow range of altitudes. Most species live near open, rocky areas. Rocks offer them cover from predators, plentiful prey (e.g. rodents, lizards, insects, etc. that live amidst the rocks), and open basking areas. However, rattlesnakes can also be found in a wide variety of other habitats including prairies, marshes, deserts, and forests.[6] Rattlesnakes prefer a temperature range between 80 and 90°F (26-32°C), but can survive temperatures below freezing, recovering from brief exposure to temperatures as low as 4°F (-16°C), and surviving for several days in temperatures as low as 37°F (3°C).[7]

Evidence has been collected to show the most probable ancestral area of rattlesnakes is the Sierra Madre Occidental region in Mexico. This same study found the most probable vegetation or habitat of the ancestral area to be pine-oak forests.[8] Feeding habits play an important ecological role by limiting the size of rodent populations, which prevents crop damage and stabilizes ecosystems.[9]

Prey[edit]

Rattlesnakes consume mice, rats, small birds and other small animals.[10]

Rattlesnakes lie in wait for their prey, or hunt for it in holes.[11] The prey are killed quickly with a venomous bite as opposed to constricting. If the bitten prey moves away before dying, the rattlesnake can follow it by its scent.[12][13] When it locates the fallen prey, it checks for signs of life by prodding with its snout, flicking its tongue, and using its sense of smell. Once the prey has become incapacitated, the rattlesnake locates its head by odors emitted from the mouth. The prey is then ingested head-first, which allows wings and limbs to fold at the joints in a manner which minimizes the girth of the meal.[14] The gastric fluids of rattlesnakes are extremely powerful, allowing for the digestion of bone as well as flesh. Optimal digestion occurs when the snake maintains a body temperature of between 80 and 85°F (25-29°C). If the prey is small, the rattlesnake will often continue hunting. If it was an adequate meal, the snake will find a warm, safe location in which to coil up and rest until the prey is digested.[15]

Rattlesnakes are believed to require at least their own body weight in water annually in order to remain hydrated. The method in which they drink depends on the water source. In larger bodies of water (streams, ponds, etc.) they will submerge their head and ingest water by opening and closing their jaw, which sucks in water. If drinking dew, or drinking from small puddles, they will sip the liquid either by capillary action or by flattening and flooding the lower jaw.[16]

Predators[edit]

Newborn rattlesnakes are heavily preyed upon by a variety of species including ravens, crows, roadrunners, raccoons, opossums, skunks, coyotes and weasels, whipsnakes, kingsnakes, and racers. Neonates of the smaller Crotaline species are frequently killed and eaten by small predatory birds such as jays, kingfishers, and shrikes. Some species of ants in the genus Formica are known to prey upon neonates, and it is very likely that Solenopsis invicta (fire ants) do as well. On occasion, hungry adult rattlesnakes will cannibalize neonates. The small proportion (often as few as 20%) of rattlesnakes that make it to their second year are heavily preyed upon by a variety of larger predators including coyotes, eagles, hawks, owls, feral pigs, badgers, indigo snakes, and kingsnakes.[17]

The common kingsnake (Lampropeltis getula), a constrictor, is immune to the venom of rattlesnakes and other vipers, and rattlesnakes form part of its natural diet. Rattlesnakes sense kingsnakes' presence by their odor.[18] When they realize a kingsnake is nearby, they begin enacting a set of defensive postures known as "body bridging". Unlike its normal erect and coiled defensive-striking posture, the rattlesnake keeps its head low to the ground in an attempt to prevent the kingsnake from gaining a hold of it (the head being the first part of the rattlesnake that is ingested). The rattlesnake jerks its body about, while bridging its back upwards, forming an elevated coil which faces the kingsnake. The elevated coil is used to strike the attacker, and is also used to shield the head from the kingsnake.[19]

Anatomy[edit]

Sensory organs[edit]

Timber rattlesnake (Crotalus horridus) with clearly visible facial pits

Like all pit vipers, rattlesnakes have two organs that can sense radiation: their eyes, and a set of heat-sensing "pits" on their face that enable them to locate prey and move towards it, based on the prey's thermal radiation signature. These pits have a relatively short effective range of approximately 1 ft, but nevertheless give the rattlesnake a distinctive advantage in hunting for warm-blooded creatures at night.[20][21]

Heat Sensing Pits[edit]

Aside from this pair of simple eyes, rattlesnakes are able to detect thermal radiation emitted by warm-blooded organisms in their environment. As mentioned previously, these pit organs are positioned between the eye and nostril on both sides of the rattlesnake’s head. Functioning optically like a pinhole camera eye, thermal radiation, in the form of infrared wavelength light, enters passes through the opening of the pit and strikes the pit membrane located in the back wall warming this part of the organ. Due to the extremely high density of these heat sensitive receptors innervating this membrane, the rattlesnake can detect temperature changes of 0.003°C or less in its immediate surroundings. Infrared cues from these receptors are transmitted to the brain via the trigeminal nerve where they are used to create thermal maps of the snake’s surroundings. Due to the small sizes of the pit openings, typically these thermals images are low in resolution and contrast. Nevertheless, rattlesnakes superimpose visual images created from information from the eyes with these thermal images from the pit organs in order to more accurately visualize their surroundings in low levels of light. Research conducted recently on the molecular mechanism of this ability suggests that the temperature sensitivity of these pit organs is closely linked to the activity of TRPA1 (transient receptor potential ankyrin 1), a temperature sensitive ion channel saturated in the pit membrane. This ability to detect infrared radiation in their surrounds enhances the rattlesnake’s ability to hunt prey and identify predators during periods of low-level light.

Eyes[edit]

Rattlesnake eyes, which contain a large number of rod cells, are well adapted to nocturnal use.[22][23] However, rattlesnakes are not exclusively nocturnal, and their vision is more acute during daylight conditions.[23] Rattlesnakes also possess cone cells, which means that they are capable of some form of color vision. The rattlesnake eye lacks a fovea making it impossible for them to see sharply defined images. Instead, they mostly rely on the perception of movement.[22] Rattlesnake eyes are capable of horizontal rotation, but they do not appear to move their eyeballs to follow moving objects.[23]

Smell[edit]

Rattlesnakes have an exceptionally keen sense of smell. They can sense olfactory stimuli both through their nostrils, and by flicking their tongue, which carries scent-bearing particles to the Jacobson's organ in the roof of their mouth.[24][25]

Auditory system[edit]

Like all snakes, rattlesnakes do not have external ear openings, and the structures of their middle ear is not as highly specialized as those of other vertebrates, such as mammals. Thus their sense of sound is not very effective. However, they are capable of sensing vibrations in the ground, passed via the skeleton to the auditory nerve.[22]

Fangs[edit]

Rattlesnake skull

Rattlesnake fangs are connected by venom ducts to large poison glands near the outer edge of the upper jaw, towards the rear of the head. When the rattlesnake bites, muscles on the sides of the venom glands contract, which squeezes the venom through the ducts and into the fangs. When the fangs are not in use, they remain folded against the palate.[26][27]

Rattlesnakes are born with fully functioning fangs and venom and are capable of killing prey at birth.[17][28] Adult rattlesnakes shed their fangs every 6–10 weeks. At least 3 pairs of replacement fangs lie behind the functional pair.[29]

Venom[edit]

Most species of rattlesnakes can control how much venom to inject and have hemotoxic venom, destroying tissue, causing necrosis and coagulopathy (disrupted blood clotting).[30] In the U.S., the Tiger rattlesnake (C. tigris) and some varieties of the Mojave rattlesnake (C. scutulatus) also have a presynaptic neurotoxic venom component known as Mojave Type A toxin, which can cause severe paralysis.[30][31][32] Although it has a comparatively low venom yield,[33] the venom toxicity of C. tigris is considered to be the highest of all rattlesnake venoms, and the highest of all snakes in the Western Hemisphere based on LD50 studies conducted on laboratory mice. C. scutulatus is also widely regarded as producing one of the most toxic snake venoms in the Americas, based on LD50 studies in laboratory mice.[34]

Rattlesnake venom is a mixture of 5-15 enzymes, various metal ions, biogenic amines, lipids, free amino acids, proteins, and polypeptides. It contains components designed to immobilize and disable the prey, as well as digestive enzymes which break down tissue to prepare for later ingestion.[25][29] The venom is very stable, and retains its toxicity for many years in storage.[25]

Older snakes possess more potent venom, and larger snakes are frequently capable of storing larger volumes of it.[35]

Rattle[edit]

Rattlesnake rattle
Drawing of the rattle
The characteristic sound of a rattlesnake -- the rattle shaking, as a result of extremely rapid tail movement.

Problems playing this file? See media help.

According to Rubio, "The most widely accepted hypothesis for the evolution of the rattle is that the rattle is a warning device for predatory animals that might be a threat to the rattlesnake. It produces a signal to drive them away."[36]

The rattle is composed of a series of hollow, interlocked segments made of keratin, which are created by modifying the scales that cover the tip of the tail. The contraction of special "shaker" muscles in the tail causes these segments to vibrate against one another, making the rattling noise (which is amplified because the segments are hollow).[1][37][38] The muscles that cause the rattle to shake are some of the fastest known, firing 50 times per second on average, sustained for up to three hours.[39]

At birth, a “pre-button” is present at the tip of the snake's tail; it is replaced by the “button” several days later when the first skin is shed. However, no sound can be made by the rattle until a second segment is added when the skin is shed again.[40] A new rattle segment is added each time the snake sheds its skin, and the snake may shed its skin several times a year, depending on food supply and growth rate.

Rattlesnakes travel with their rattles held up to protect them from damage, but in spite of this precaution, their day-to-day activities in the wild still cause them to regularly break off end segments. Because of this, the age of a rattlesnake is not related to the number of rattles on its tail.[1][38][39]

One of the differentiating features of males and females is that males have thicker and longer tails (because they contain the inverted hemipenes). Also, the tails of males taper gradually from the body, whereas the tails of females narrow abruptly at the vent.[41]

Skin and circulation[edit]

Rattlesnake skin is composed of a set of overlapping scales which cover the entire body, providing protection from a variety of threats including dehydration and physical trauma.[42] The typical rattlesnake--genus Crotalus--have the top of the head covered with small scales, except, with a few species, a few crowded plates directly over the snout.[43] The skin of snakes is highly sensitive to contact, tension, and pressure; and they are capable of feeling pain.[44]

An important function of the skin is the sensation of changes in air temperature, which can guide the snakes towards warm basking/shelter locations.[45] All snakes are ectotherms. In order to maintain a stable body temperature, they exchange heat with their external environment. Snakes often move into open, sunny areas to absorb heat from the sun and warmed earth - a behavior known as basking. Nerves in the skin regulate the flow of blood into the veins near the surface. Rubio states that "The skin's acceptance of radiant energy, its ability to determine the temperature, and the snake's ability to move toward or away from one temperature gradient to another are among the most important behavioral actions in its daily life."[44]

The skin of rattlesnakes is intricately patterned in a manner that camouflages them from their predators.[45][46] Rattlesnakes do not generally have bright or showy colors (reds, yellows, blues, etc.), instead relying on subtle earth tones that resemble the surrounding environment.[47]

There are creases in the epidermal tissue which connects the scales of rattlesnakes. When ingesting large prey, these creases can unfold, allowing the skin to expand to envelop a much greater volume. It appears that the skin is tightly stretching to accommodate the meal, but in reality, the skin is simply smoothing out from its creased state and is not under very high tension.[48]

Reproduction[edit]

Two male Northern Pacific Rattlenakes (Crotalus oreganus oreganus) engage in a "combat dance"
Everted hemipenis of a male C. adamanteus

Most rattlesnake species mate during the summer or fall, while some species mate only in the spring, or during both the spring and fall.[49]

Females secrete small amounts of sex pheromones, which leave a trail that males follow by using their tongue and Jacobson's organ as a guide.[49] Once a receptive female has been located, the male will often spend several days following her around (a behavior that is not common outside of the mating season), frequently touching and rubbing her in an attempt to stimulate her.[50][51]

The males of some species, such as timber rattlesnakes (C. horridus), will fight each other during the mating season, in competition over females. These fights known as "combat dances" consist of the two males intertwining the anterior portion of their bodies, often with the head and neck held vertically. The larger males usually end up driving the smaller males away.[52][53]

Although many kinds of snakes and other reptiles are oviparous (lay eggs), rattlesnakes are ovoviviparous (give birth to live young after carrying eggs inside).[54] The female produces the ova ("eggs") in her ovaries, after which they pass through the body cavity and into one of her two oviducts. The ova are arranged in a continuous chain in a coiled section of the oviduct, known as the "tuba".[50] Male rattlesnakes have sexual organs known as hemipenes, located in the base of the tail. The hemipenis is retracted inside of the body when mating is not occurring. Females can store semen for months in internal recesses known as spermathecae, which permits them to mate during the fall, but not fertilize the ova until the following spring.[50] The Arizona black rattlesnake (C. oreganus cerberus), has been observed to exhibit complex social behavior reminiscent of that in mammals. Females often remain with their young in nests for several weeks, and mothers have been observed cooperatively parenting their broods.[55]

Rattlesnakes generally take several years to mature, and females usually reproduce only once every three years.[56]

Hibernation[edit]

Some rattlesnake species hibernate in the colder winter months. They often gather together for hibernation in very large numbers (sometimes over 1,000 snakes), huddling together inside underground "rattlesnake dens" or hibernacula.[57][58] Rattlesnakes will regularly share their winter burrows with a wide variety of other species (such as turtles, small mammals, invertebrates, and other types of snakes).[57]

Rattlesnakes often return to the same den, year after year, sometimes traveling several miles to get there. It is not known exactly how the rattlesnakes find their way back to the dens each year, but it has been hypothesized that they use a combination of pheromone trails and visual cues (e.g. topography, celestial navigation, and solar orientation).[59]

Species with long periods of hibernation tend to have much lower reproductive rates than those with shorter hibernation periods, or those that don't hibernate at all. Female timber rattlesnakes in high peaks in the Appalachian Mountains of New England reproduce every three years on average; the lance-headed rattlesnake (C. polystictus), native to the warm climate of Mexico, reproduces annually.[60]

Like most other snakes, rattlesnakes aestivate during very hot or dry periods, which is why they are very rarely seen during the hottest and driest months of summer.[61]

Conservation status[edit]

Rattlesnakes tend to avoid developed areas, preferring undisturbed, natural habitats. Rapid habitat destruction by humans, mass killings during events like rattlesnake round-ups, and deliberate extermination campaigns all pose severe threats to rattlesnake populations in many areas. Several species of rattlesnakes, such as the timber rattlesnake, massasauga, and canebrake rattlesnake are listed as threatened or endangered in many U.S. states.[62]

A very large number of rattlesnakes die from being run over by cars.[17]

In more heavily populated and trafficked areas, there have been increasing reports of rattlesnakes that do not rattle. This phenomenon is commonly attributed to selective pressure by humans, who often kill the snakes when they are discovered.[63] Non-rattling snakes are more likely to go unnoticed, and therefore survive to reproduce offspring who, like themselves, are less likely to rattle.

Safety and first aid[edit]

A rattlesnake warning sign in California

Rattlesnakes are the leading cause of snakebite injuries in North America and a significant cause in Central and South America.[32][64]

Avoiding bites[edit]

Rattlesnakes tend to avoid wide open spaces where they cannot hide from predators and will generally avoid humans if they are aware of their approach.[65] Rattlesnakes rarely bite unless they feel threatened or provoked. A majority of victims (~ 72%[66]) are males, often young and intoxicated. Approximately half of bites occur in cases where the victim saw the snake yet made no effort to move away.[25]

Hikers and campers should avoid contact with rattlesnakes by remaining observant and not approaching the animals. Hikers should be particularly careful when negotiating fallen logs or boulders and when near rocky outcroppings and ledges where rattlesnakes may be hiding or sunning themselves. Snakes will occasionally sun themselves in the middle of an open trail so such areas are not the only places where they are encountered. Hikers are advised to keep their distance when encountering a rattlesnake on a trail and allow the snake room to retreat.[citation needed]

Caution is advised even when snakes are believed to be dead; rattlesnake heads can see, flick the tongue, and inflict venomous bites for up to an hour after being severed from the body.[67][68]

Effect of bites on humans[edit]

Rattlesnake bite injury

Rattlesnake bites are rarely fatal to humans if treated promptly.[30] An estimated 7,000 to 8,000 people are bitten by venomous snakes in the United States each year, with approximately five deaths.[69] The most important factor in survival following a severe envenomation is the amount of time elapsed between the bite and treatment. Most deaths occur between 6 and 48 hours after the bite. If antivenom treatment is given within 2 hours of the bite, however, the probability of recovery is greater than 99%.[70]

When a bite occurs the amount of venom injected is under voluntary control by the snake. The amount released depends on a variety of factors including the condition of the snake (e.g. having long, healthy fangs and a full venom sack) and its temperament (an angry, hungry snake that has just been stepped on vs. a satiated snake that was merely surprised by walking near it).[71] Approximately 20% of bites result in no envenomation at all. A lack of burning pain and edema 38 in (1 cm) away from the fang marks after 1 hour suggests that either no or minimal envenomation occurred. A lack of edema or erythema in the area of the bite after 8 hours indicates a lack of envenomation for most rattlesnake bites.[72]

Common symptoms include swelling, severe pain, tingling, weakness, anxiety, nausea and vomiting, hemorrhaging, perspiration, and eventually heart failure.[71][73] Local pain following envenomation is often intense, increasing with the ensuing edema.[71] Children generally experience more severe symptoms because they receive a larger amount of venom per unit of body mass.[72]

Emergency response[edit]

Data on the effectiveness of first aid techniques for rattlesnake bites is limited.[74] General recommendations for first aid in the field are:

  1. Remain calm and retreat from the snake at least 15 feet. Arrange to have the victim transported to a medical facility as soon as possible.[74]
  2. Remove restrictive clothing items (rings, bracelets, watches, buttoned shirts, etc.) from the victim.[74][75]
  3. Splint or otherwise immobilize any bitten limbs and keep them below heart level.[30][35][74][76] If (and only if) the victim is more than 1 hour away from a medical facility place a lightly constricting band (that admits one finger beneath it) above the bitten area to prevent the systemic spread of the venom.[74]
  4. Keep victims calm; put them at rest; keep them warm and give them comfort and reassurance (which will lower their heart rate, slowing the spread of the venom). Keeping a victim's heart rate down, however, should never interfere with getting him or her to a medical facility.[74]
  5. If the snake is still present or nearby, try to get an accurate description by using a camera or remembering certain physical traits such as color, pattern, or length. This can help ensure the proper anti venom is administered. However, one should never put themselves at risk of being bitten to obtain this description.

In no case should tourniquets be used, nor should any incisions or suction be applied to the wound.[74][76]

Antivenin[edit]

Antivenin, often referred to as antivenom, is commonly used to treat the effects of local and systemic pit viper envenomations.[77] The first step in the production of crotaline antivenin is collecting ("milking") the venom of a live rattlesnake—usually from the Western Diamondback (Crotalus atrox), Eastern Diamondback (Crotalus adamanteus), South American rattlesnake (Crotalus durissis terrificus), or fer-de-lance (Bothrops atrox). The extracted venom is then diluted and injected into horses, goats, or sheep, whose immune systems produce antibodies that protect from the toxic effects of the venom. These antibodies accumulate in the blood, which is then extracted and centrifuged to separate the red blood cells. The resulting serum is purified into a lyophilized powder, which is packaged for distribution and later use by human patients.[78][79]

Because antivenin is derived from animal antibodies, people generally display an allergic response during infusion, known as serum sickness.[80][81]

Veterinary care[edit]

In the United States, more than 15,000 domesticated animals are bitten by snakes each year. Rattlesnake envenomations account for 80% of the deadly incidents.[82]

Dogs are most commonly bitten on the front legs and head. Horses generally receive bites on the muzzle, and cattle on their tongues and muzzles. If a domesticated animal is bitten, the hair around the bite should be removed, so that the wound can be clearly seen. The crotaline Fab antivenin has been shown to be effective in the treatment of canine rattlesnake bites. Symptoms include swelling, slight bleeding, sensitivity, shaking, and anxiety.[35]

In human culture[edit]

Spirituality[edit]

Stone sculptures of feathered serpents on display at the National Museum of Anthropology in Mexico City.

Indigenous Americans[edit]

Aztec paintings, Central American temples, and the great burial mounds in the southeastern U.S. are frequently adorned with depictions of rattlesnakes, often within the symbols and emblems of the most powerful deities.[83]

The Feathered Serpent deity of Olmec, Aztec and Mayan cultures was depicted as having the combined features of the quetzal bird and rattlesnake.[84] The Ancient Maya considered the rattlesnake to be a "vision serpent" that acted as a conduit to the "otherworld".[85]

Rattlesnakes are a key element in Aztec mythology and were widely represented in Aztec art including sculptures, jewelry, architectural elements, etc.

Christian snake handling sects[edit]

Snake handling at the Pentecostal Church of God in the town of Lejunior, Harlan County, Kentucky September 15, 1946

Members of some Christian sects in the southern United States are regularly bitten while participating in "snake handling" rituals. Snake handling is when people hold venomous snakes, unprotected, as part of a religious service inspired by a literal interpretation of the Bible verse Mark 16:17-18 which reads "In my name ... they will pick up snakes with their hands".[25][86]

As food[edit]

Journalist Alistair Cooke claims that rattlesnake tastes "just like chicken, only tougher."[87] Others have compared the flavor to a wide range of other meats, including veal, frog, tortoise, quail, fish, rabbit, and even canned tuna.[88]

See also[edit]

References[edit]

  1. ^ a b c Price, Andrew H. (2009). Venomous Snakes of Texas: A Field Guide. University of Texas Press. pp. 38–39. ISBN 978-0-292-71967-5. 
  2. ^ Barceloux, 2008: p. 1026
  3. ^ Willis Lamm. "RATTLESNAKE!". 
  4. ^ Kini, R. Manjunatha et al., ed. (2011). Toxins and Hemostasis. Springer 2011. p. 99. ISBN 978-90-481-9294-6. 
  5. ^ Fox, William Sherwood (1988). The Bruce beckons: the story of Lake Huron's great peninsula. University of Toronto Press. p. 122. ISBN 978-0-8020-6007-5. 
  6. ^ Rubio, 1998: p. 24
  7. ^ Rubio, 1998: p. 71
  8. ^ Place, Arron J; Abramson, Charles I. (2004). "A Quantitative Analysis of the Ancestral Area of Rattlesnakes". Journal of Herpetology 38 (1): 151–156. doi:10.1670/103-03N. 
  9. ^ Rubio, 1998: pp. 161, 163
  10. ^ Klauber, 1997: p. 612
  11. ^ Klauber, 1997: p. 387
  12. ^ Klauber, 1997: p. 834
  13. ^ Parker, M. Rockwell & Kardong, Kenneth V. (2005). "Rattlesnakes can use airborne cues during post-strike prey relocation". In Mason, Robert T. et al. Chemical signals in vertebrates 10. Springer. p. 397. ISBN 978-0-387-25159-2. 
  14. ^ Rubio, 1998: p. 81
  15. ^ Rubio, 1998: p. 83
  16. ^ Rubio, 1998: p. 87
  17. ^ a b c Rubio, 1998: p. 120
  18. ^ Rubio, 1998: p. 59 -- Knowledge that rattlesnakes are so afraid of kingsnakes' odor led to the development of effective synthetic rattlesnake repellents.
  19. ^ Rubio, 1998: p. 59 -- This behavior is believed to be unique to crotaline species.
  20. ^ Klauber, 1997: pp.401-402
  21. ^ Furman, 2007: p. 8
  22. ^ a b c Rubio, 1998: p. 67
  23. ^ a b c Klauber, 1997: pp. 384-389
  24. ^ Furman, 2007: p. 9
  25. ^ a b c d e Cetaruk, Edward W. (2005). "Rattlesnakes and Other Crotalids". In Brent, Jeffrey. Critical care toxicology: diagnosis and management of the critically poisoned patient. Elsevier Health Sciences. p. 1075. ISBN 978-0-8151-4387-1. 
  26. ^ Vitts, Laurie J. (1999). "Rattlesnake". In Mares, Michael A. et al. Encyclopedia of deserts. University of Oklahoma Press. p. 468. ISBN 978-0-8061-3146-7. 
  27. ^ Klauber, 1997: p. 773
  28. ^ See also Klauber, 1997: p. 829
  29. ^ a b Barceloux, 2008: p. 1028
  30. ^ a b c d Schoenherr, Allan A. (1995). A Natural History of California. University of California Press. p. 510. ISBN 978-0-520-06922-0. 
  31. ^ Lessenger, James E., ed. (2006). Agricultural medicine: a practical guide. Birkhäuser. p. 447. ISBN 978-0-387-25425-8. 
  32. ^ a b Luch, Andreas, ed. (2010). Molecular, Clinical and Environmental Toxicology 2. Springer. p. 267. ISBN 978-3-7643-8337-4. 
  33. ^ Weinstein and Smith (1990)
  34. ^ Glenn, J.L., R.C.Straight. 1982. The rattlesnakes and their venom yield and lethal toxicity. In: Tu, A. (ed) Rattlesnake Venoms, Their Actions and Treatment. New York: Marcel Dekker, Inc.
  35. ^ a b c Gupta, Ramesh Chandra, ed. (2007). Veterinary toxicology: basic and clinical principles. Academic Press. pp. 800–801. ISBN 978-0-12-370467-2. 
  36. ^ Rubio, 1998: p. 56
  37. ^ Burton, Maurice & Burton, Robert, ed. (1970). "Rattlesnake". The international wildlife encyclopedia, Volume 1. Marshall Cavendish. p. 2119. ISBN 978-0-7614-7266-7. 
  38. ^ a b Fergus, Charles (2003). Wildlife of Virginia and Maryland and Washington, Part 3. Stackpole Books. p. 460. ISBN 978-0-8117-2821-8. 
  39. ^ a b Graham, Karen S. (2001). "Rattlesnake". In Bell, Catharine E. Encyclopedia of the world's zoos, Volume 3. Taylor & Francis. p. 1040. ISBN 978-1-57958-174-9. 
  40. ^ American International Rattlesnake Museum: Rattles, (Accessed: May 4, 2012)
  41. ^ Klauber, 1997: p. 690
  42. ^ Rubio, 1998: p. 48
  43. ^ Ditmars, Raymond L. "Reptiles of the World" The MacMillan Company, New York, 1936, p. 255
  44. ^ a b Rubio, 1998: p. 69
  45. ^ a b Adams, Clark E. & Thomas, John K. (2008). Texas Rattlesnake Roundups. Texas A&M University Press. p. 2. ISBN 978-1-60344-035-6. 
  46. ^ Rubio, 1998: p. 38
  47. ^ Rubio, 1998: p. 39
  48. ^ Rubio, 1998: p. 50
  49. ^ a b Rubio, 1998: p. 106
  50. ^ a b c Rubio, 1998: p. 110
  51. ^ Klauber, 1997: p. 702
  52. ^ Furman, 2007: p. 32
  53. ^ Rubio, 1998: p. 109
  54. ^ http://www.nwf.org/Wildlife/Wildlife-Library/Amphibians-Reptiles-and-Fish/Rattlesnakes.aspx
  55. ^ Amerello, Melissa; Smith, Jeffrey; and Slone, John. 2011. Family values: Maternal care in rattlesnakes is more than mere attendance. Nature Precedings. <http://dx.doi.org/10.1038/npre.2011.6671.1>.
  56. ^ Hammerson, Geoffrey A. (2004). Connecticut wildlife: biodiversity, natural history, and conservation. UPNE. p. 328. ISBN 978-1-58465-369-1. 
  57. ^ a b Rubio, 1998: p. 96
  58. ^ Klauber, 1997: p. 573
  59. ^ Rubio, 1998: pp. 96-97
  60. ^ Furman, 2005: p. 33
  61. ^ Rubio, 1998: p. 100
  62. ^ Rubio, 1998: pp. 200-203
  63. ^ "Rattlesnake danger grows as more serpents strike without warning". The San Francisco Chronicle. June 24, 2011. 
  64. ^ Mackessy, Stephen P., ed. (2009). Handbook of venoms and toxins of reptiles. CRC Press. p. 476. ISBN 978-0-8493-9165-1. 
  65. ^ Phillips, Steven J. et al., ed. (2009). A natural history of the Sonoran Desert. University of California Press. p. 577. ISBN 978-0-520-21980-9. 
  66. ^ O'Neil ME, Mack KA, Gilchrist J, Wozniak EJ (2007). "Snakebite injuries treated in United States emergency departments, 2001-2004". Wilderness Environ Med 18 (4): 281–7. doi:10.1580/06-WEME-OR-080R1.1. PMID 18076294. 
  67. ^ Werler, John E. & Dixon, James Ray, ed. (2000). Texas snakes: identification, distribution, and natural history. University of Texas Press. p. 3. ISBN 978-0-292-79130-5. 
  68. ^ Barceloux, 2008: p. 1027
  69. ^ Henkel, John. "For Goodness Snakes! Treating and Preventing Venomous Bites". Reptiles. USDA / emergency response. Retrieved 2009-06-15. 
  70. ^ Rubio, 1998: p. 143
  71. ^ a b c Fleisher, Gary R. & Ludwig, Stephen, ed. (2010). Textbook of Pediatric Emergency Medicine (6th ed.). Lippincott Williams & Wilkins. p. 1548. ISBN 978-1-60547-159-4. 
  72. ^ a b Barceloux, 2008: p. 1030
  73. ^ Klauber, 1997: p. 859
  74. ^ a b c d e f g Barceloux, 2008: p. 1033
  75. ^ Tibballs, James et al. (2010). "Envenomations". In Wheeler, David S. et al. Pediatric Critical Care Medicine: Basic Science and Clinical Evidence (6th ed.). Springer. p. 1693. ISBN 978-1-84628-463-2. 
  76. ^ a b Taylor, Robert B., ed. (2002). Manual of family practice (2nd ed.). Lippincott Williams & Wilkins. p. 632. ISBN 978-0-7817-2652-8. 
  77. ^ Goldfrank, Lewis R., ed. (2006). Goldfrank's toxicologic emergencies (8th ed.). McGraw-Hill Professional. p. 1657. ISBN 978-0-07-147914-1. 
  78. ^ Barceloux, 2008: p. 1036
  79. ^ Adams, Clark E. & Thomas, John K. (2008). Texas Rattlesnake Roundups. Texas A&M University Press. p. 23. ISBN 978-1-60344-035-6. 
  80. ^ Meier, Jürg & White, Julian, ed. (1995). Handbook of clinical toxicology of animal venoms and poisons, Volume 236. CRC Press. p. 639. ISBN 978-0-8493-4489-3. 
  81. ^ Rubio, 1998: p. 145
  82. ^ Slatter, Douglas H. (2002). Textbook of small animal surgery. Elsevier Health Sciences. p. 267. ISBN 978-0-7216-8607-3. 
  83. ^ Browman, David L. & Williams, Stephen (2002). New perspectives on the origins of Americanist archaeology. University of Alabama Press. p. 99. ISBN 978-0-8173-1128-5. 
  84. ^ Read, Kay A. & González, Jason J., ed. (2002). "Feathered Serpents". Mesoamerican Mythology: A Guide to the Gods, Heroes, Rituals, and Beliefs of Mexico and Central America. Oxford University Press. p. 180. ISBN 978-0-19-514909-8. 
  85. ^ Foster, Lynn V. & Mathews, Peter, ed. (2005). "Maya Geography and Mythology". Handbook to life in the ancient Maya world. Oxford University Press. p. 91. ISBN 978-0-19-518363-4. 
  86. ^ For a more detailed study of snake handling sects, see Kimbrough, David L. (2002). Taking up serpents: snake handlers of eastern Kentucky. Mercer University Press. ISBN 978-0-86554-798-8. 
  87. ^ Cooke, Alistair (1980). The Americans: fifty talks on our life and times. Knopf. p. 183. ISBN 978-0-394-50364-6. OCLC 5311048. "To the goggling unbeliever Texans say—as people always say about their mangier dishes—'but it's just like chicken, only tenderer.' Rattlesnake is, in fact, just like chicken, only tougher." 
  88. ^ Klauber, 1997: p. 1055

Further reading[edit]

External links[edit]