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amphibians, frogs, salamanders, caecilians, newts, mudpuppies, waterdogs, axolotls, sirens, amphiuma, biology, physiology, genetics, distribution maps, zoology, conservation, preservation, biodiversity
Article 0011 Toxicity & Defense Methods of Amphibians

Introduction


Amphibians are a diverse group of animals that have evolved numerous different adaptations to their environments. Some of the most interesting of these adaptations are the varying methods of defense portrayed by frogs, salamanders, and caecilians. Because they inhabit such a diverse range of environments, many species have developed unique and amazing methods of warding off intruders and predators, including the possession of toxic chemicals (toxicity), foul-tasting secretions, startling eyes, ribs that pierce the skin and act as tiny swords, mimicry of predatory animals, inflating the body with air (bloating), and more. This article is meant to introduce the different defense methods of amphibians, with emphasis on the toxicity of newts of the family Salamandridae (order: Caudata).

Although many amphibian species are toxic and/or noxious, they are passive and do not appreciate disturbances. However, when living in a predator-prey environment, attacks and disturbances from curious or hungry creatures are inevitable. In response, amphibians have evolved several different methods of warding off potential predators and attackers. This section is a general introduction to the varying methods of amphibian defense mechanisms, separated into two section covering Anuran and Caudate species.

Amphibian Skin Secretions: Mucus and Toxins


Most amphibians, if not all, often secrete fluids from epidermal glands, in the form of protective mucus or toxins. These glands are distributed throughout the skin of the amphibian. Poison glands, or granular glands, are larger than mucous glands. Both types of glands are alveoliform, meaning they extend through the epidermis, ending in a circular pore. Some amphibians, such as toads, newts, and salamanders, possess additional glands with high concentrations of toxins, called paratoid glands. Paratoid glands are visible to varying degrees, and are typically found behind the head. Some Salamandrids also possess paratoid glands, often referred to as warts, in dorsolateral rows along both sides of the spine. Paratoid glands are usually ruptured when pressure is applied, however, some amphibians are capable of spraying their toxins using muscle contractions.

Protective mucus coats help keep harmful microorganisms off the delicate, naked skin, as well as aid in moisture retention. Without the protective mucus, many species would be highly susceptible to desiccation and/or microbial attack. When ill, many amphibian species develop problems with glandular function, and may take on a pasty, dry appearance due to lack of a sufficient mucus coat. Some species, such as the giant salamanders (Caudata: Cryptobranchidae), can use a thick mucous coat as a defense and escape method. This type of mucous is very thick, painstaking to wash off, and make it difficult to physically handle such individuals. 

Many amphibian species are also toxic, such as the newts (Caudata: Salamandridae) and poison frogs (Anura: Dendrobatidae). Toxic amphibians still possess a mucous coat to ward off microbial attacks and prevent desiccation, however, they possess additional glands that produce toxins. Newts, for example, possess a chemical called Tarichatoxin, which is a highly effective defense against nearly every predator in the wild. The presences of toxins are usually accompanied by bright coloration, aposematic coloration, which is displayed as a warning sign when such an individual is disturbed. Some species, such as the Dendrobatids, are wholly covered in aposematic colors, allowing them to forage during the daylight, undaunted by birds, snakes, rodents, or other potential predators. Amphibian toxins have evolved independently in different amphibian groups, and so there are several different alkaloids that have been isolated in different species. Aposematic colors and the possession of toxins are still not enough for most species. Many amphibians have additional measures of defense in addition to toxins. These can range from cryptic coloration, aggressive behavior, mimicry and more.

Caudate Defense Methods


 
Toxicity & Aposematic Coloration
Tylototriton shanjing © Paolo Mazzei, http://www.herp.it Many caudates are toxic, and can cause severe damage or death to a potential predator in the wild if consumed. Those that are not toxic may have noxious secretions, or foul tasting and smelling mucous. Some caudates are inconspicuously colored, dorsally, to blend into their environment, however, some species may possess aposematic coloration (warning colors) on the dorsum and/or venter. The photo to the left shows Tylototriton shanjing, who possesses vivid coloration on the dorsum, tail, head, and feet, with a black venter. T. shanjing also possess several "warts" along the dorsum, which are actually paratoid glands. In most species, toxic secretions are administered when harassed, in conjunction with particular defensive maneuvers, outlined below. 

Immobilization & Cryptic Coloration
Desmognathus marmoratus © Rich Glor Immobilization is usually the initial response to a potential predator by most caudates. Immobilization consists of cessation of all movement and flattening of the body so as to blend into the immediate environment. If immobilization is not effective, a defense posture may be exhibited (unken reflex), along with toxic secretions, tail-lashing, biting, writhing, flipping over, etc., depending on the species. Because of their slow metabolism, caudates would benefit more from not being noticed, rather than having to make a quick get away (like frogs). This is why it's rather easy to find terrestrial caudates in the wild; they don't put much effort into running from you! Aquatics, on the other hand, are very agile and swift in the water and can dart out of harms way with little effort. Darting is usually the first and only defensive maneuver of aquatics, although if cornered, more aggressive species, such as amphiuma, hellbenders, and mudpuppies, may inflict painful bites.

To reduce the risk of detection, many species have evolved coloration that closely resembles their natural habitat. The photo at right shows a rather effectively camouflaged Desmognathus marmoratus.

Rhyacotriton olympicus - unken reflex. Photo © 2002 William FlaxingtonDefensive Posturing, the Unken Reflex, & Aposematic Coloration
 Many newts and salamanders are inconspicuously colored dorsally to blend into their habitat (cryptic coloration), but possess  brightly colored bellies to warn of their toxicity (Cynops, Triturus, Paramesotriton, Rhyacotriton etc.). When such species are adequately disturbed, they will assume a defensive posture that shows the Taricha torosa sierrase - mild defensize posturing. Photo © 2002 William Flaxingtonbrightly colored belly and underside of tail in an attempt to clarify their toxicity. A common defensive posture of the family Salamandridae consists of flattening the abdomen, and curling the tail and head back to show the bright belly. The salamander may assume a circular shape, semi-circular shape, or simply raise the head to display the bright colors. This posturing is a natural reflex stimulated by persistent disturbance, called the unken reflex. The unken reflex is typical of the genera Taricha, Triturus, Cynops, Rhyacotriton, and more. While in position, toxic secretions are usually administered from paratoid glands. In most species, the body becomes extremely tense when in the unken position.

Disruptive Coloration
Ensatina eschscholtzii xanthoptica juvenile . Photo © Jessica J. MillerSome species may be colored in a manner that breaks up the silhouette of the animal on certain backgrounds. Such coloration often looks bold and conspicuous to the human eye, especially when viewed outside of the natural environment, but is actually a rather effective method of evading or confusing predators. Disruptive coloration is found in many animals, including fish and mammals, and may be expressed as dark bands across the eyes or stripes and blotches covering strategic body areas. In Salamanders, disruptive coloration is often displayed as contrasting coloration about the proximal portions of the limbs. This essentially breaks up the silhouette by disguising the presence of the limbs, which may result in the salamander going unnoticed by potential predators. The photo at righ shows such coloration about the limbs in a juvenile Ensatina eschscholtzii xanthoptica.

 Ambystoma gracile - Photo © 2002 William Flaxington"Head Butting
Other caudates, such as some Ambystomids, portray a "head-butting" technique. This consists of holding the body high off the ground, bending the head down to portray the paratoid glands at the back of the head, and swinging the back of the head toward the predator. This ensures that any potential predators first taste will be a mouthful of poison. Head butting is usually displayed in large species, possessing very large paratoid glands, and is typically enacted in conjunction with the secretion of toxic fluids. 

In the photo at right, of Ambystoma gracile, notice the large glands behind the head. This species does not possess aposematic coloration, like some other poisonous species, but usually doesn't have a problem clarifying its toxicity to potential predators. Head butting is usually a secondary defense method, performed if immobilization is unsuccessful. Ambystoma gracile, and other species, will secrete a milky, toxic substance from the glands during the head-butting display, and sometimes lash the tail, as well.

Ensatina klabauri - regenerating tail. Photo © Suzanne L. Collins, http://www.cnah.orgTail Lashing, Tail Undulating, & Tail Autotomy
Tail lashing is exhibited by caudates with toxic concentrations in the tail, and by those that can autotomize their tails. Tail lashing consists of forcefully lashing the tail back and forth in a whip-like manner. This draws the attention of the attacker to the tail, which is advantageous to the salamander for two reasons: 1) The attention is now drawn to a highly-toxic part of the salamander, and away from the head and abdomen area, 2) The tail can withstand significant damage without long term harm to, or the death of the salamander, and can be regenerated if bitten off. Some species, especially Plethodontids, do not possess tails strong enough to exhibit tail lashing, and instead undulate the tail in a vertical or semi-vertical position. Like tail lashing, tail undulating is meant to draw the attention of the attacker to the tail region, where high concentrations of toxins await. Some species can also autotomize the tail (drop if off), allowing a brief moment for escape. 

The tail can be regenerated, however this takes energy and food reserves, and also prevents the salamander from using this defense again in the near future. For these reasons, tail autotomy is usually a final effort. The in this section shows an Ensatina eschscholtzii klauberi with a partially regenerated tail. Regeneration is a costly process in this species; one that can take up to two years. Tail autotomy in species that rely heavily on energy reserves stored in the tail may have difficulty during their active time of the year, which may prevent reproduction until the tail is mostly regenerated.

Batrachoseps attenuatus. Photo © Jessica J. MillerCoiling, Rolling, & Mimicry
Some species, such as those of the genera Batrachseps and Hydromantes, will coil the body into a tight spiral when disturbed. Coiling has an added benefit for some populations of Batrachoseps attanuatus, a species that may display a dorsal stripe in such a manner that it resembles a distasteful millipede. If this does not work, B. attenuatus may begin flipping over in an abrupt and violent manner in an attempt to confuse its attacker. B. attenuatus can also autotomize its writhing tail in an attempt to distract its attacker, which may give the salamander a moment to sneak away unnoticed.

Hydromates platycephalus, and possibly other Hydromantes species, have taken coiling a setup further. They use their steeply sloped habitats as a means of escape in a coiled position. Basically, they coil up and roll away down the steep slope. This species inhabits the steep slopes of the northern Sierra Nevada, California, where rolling down a hill can provide a very quick and efficient escape, and when executed properly, leaves an attacker in a state of confusion. 

Mimicry of more toxic salamander species is observed in some species, such as Pseudotriton ruber. This bright red salamander is thought to mimic the red coloration of toxic efts of the species Notophthalmus viridescens viridescens. Western subspecies of Ensatina escscholtzii, namely the xanthoptica subspecies, are thought to be mimics of the highly toxic Taricha newts, with dark brown dorsa and yellow colored eyes and surrounding ocular areas. 

Salamandra salamandra gigliolii © Paolo Mazzei, http://www.herp.itSpecial Adaptations
The paratoid glands of most amphibian species are activated when pressure is applied, resulting in the rupture of the glands. Salamandra salamandra spp. however, are capable of spraying poison from their paratoid glands up to seven feet. This is accomplished by the contraction of muscles surrounding the paratoid glands. S. salamandra will aim the poison spray at its attacker by tilting the body in the required direction. A close-up view of a fire salamander reveals dozens of pits along the dorsum and paratoid glands. In addition to their ability to use their toxins as a direct defense when disturbed, this species displays its aposematic coloration about the entire body. The photo to the right shows Salamandra salamandra gigliolii.     

Pleurodeles waltl and Echinotriton andersonii, and possibly a few other species, have a unique defense in addition to their toxicity. These species possess paratoid glands situated dorsolaterally along the sides of body, just above the rib peaks. If threatened, these species can rupture the paratoid glands with the rib peaks when pressure is applied, such that the rib peaks break the surface of the skin. This gives the predator a mouthful of concentrated poison, and essentially injects the poison into the soft tissues of the attacker. P. waltl in particular is mostly aquatic, and so tends to use quick escapes as the primary defensive method. 


Anuran Defense Methods


 
Bufo guttatus © Paolo Mazzei, http://www.herp.itToxicity & Aposematic Coloration
Dendrobates histrionicus © Arachnokulture, http://www.pumilio.com Some anurans are toxic, and can cause severe damage or death to a potential predator in the wild. The most commonly known toxic frogs are those of the genera Dendrobates, Phyllobates, and Epipedobates, which are collectively referred to as poison frogs. There are other species, such as those of the genera Bufo, that also possess dangerous toxins capable of causing harm to potential predators. Bufids, and other species, have conspicuous paratoid (poison) glands just behind the head, capable of administering a lethal dose of poison to certain animals. Those that are not toxic may have noxious secretions, or four tasting and smelling mucous. Many anurans are inconspicuously colored, dorsally, to blend into their environment, however, the more toxic species may possess aposematic coloration on the dorsum and/or venter. Poison frogs are most noted for their full-body aposematic coloration, which allows them to lead casual, diurnal lives on the forest floors of their habitats. In most species, toxic secretions are administered when harassed, in conjunction with particular defensive maneuvers, outlined below. The photo at the left shows Bufo guttatus, whose large paratoid glands are easily detectable on the smooth dorsal side. The photo to the right shows Dendrobates histrionicus, a poison frog with a contrasting yellow and rust colored body. In general, colors of yellow, red, orange, and usually blue are aposematic colors in the wild.

Quick Getaway
Instead of immobilization, as in caudates, anurans opt for a quick getaway when harassed. The preferred course of action to a disturbance is a quick leap out of the vicinity. An abrupt leap away usually confuses potential predators long enough for the frog to make a few more leaps to a safe distance from the disturbance. This behavior is especially notable in treefrogs that leap long distances between leaves.

Bombina variegata © Paolo Mazzei, http://www.herp.itDefensive Posturing, the Unken Reflex, & Aposematic Coloration
Bombina variegata © Paolo Mazzei, http://www.herp.itLike caudates, some toxic anurans also display the unken reflex when disturbed. This is common in Bombina spp., the fire belly toads. Like caudates, the purpose of the unken reflex is to clarify the animals toxicity by displaying aposematic coloration. Fire belly toads are inconspicuously colored dorsally (cryptic coloration), but possess vivid colored bellies (aposematic coloration). Toxic secretions are usually administered from paratoid glands, and the body becomes extremely tense when in the unken position. Some species will also bloat the body, in an attempt to appear much larger. The photos at left and right show Bombina variegata displaying the unken reflex. In the photo on the left, the toad has flattened the body, and gone into a "back bend" position to show the brightly colored areas of the venter. The photo on the right shows B. variegata flipped over on its back to reveal the entirety of the yellow belly.

Hyla versicolorCryptic Coloration & Mimicry
Many amphibian species are colored in such a way so as to blend, undetected, into their immediate environment. This is called cryptic coloring, and is found in varying degrees among many amphibian species. The photo to the left shows four Hyla versicolor clinging to a tree trunk. Their varying greenish gray coloration allows them to blend nicly into the trunks, branches, and mosses of their habitat. Even when the image is enlarged, the frogs appear remarkably camouflaged. 

Some species have developed coloration and patterns to specifically mimic another organism. When disturbed, these species will flash an area of the body that resembles another animal, which is confusing to the harasser. The mimicked color pattern is usually that of the eyes of a predator, such as an owl or a snake. This type of mimicry is called Self Mimicry, and occurs when one species possess certain markings, usually eye-spots, meant to fool the predator into thinking the prey is a more dangerous animal. Although this is usually not enough to completely deter a potential predator, it may allow the amphibian a narrow opportunity to leap away while the harasser tries to figure out what's going on.

Some have gone a step further, and entirely mimic other species. Dendrobates imitator has been noted as imitating the color patterns of other sympatric Dendrobates species, with remarkable accuracy. This form of mimicry is called Müllerian Mimicry, and occurs when two toxic or noxious species mimic each others aposematic coloration. When a predator learns of the toxicity of a particular individual, it will relate the color scheme of the particular frog to the bad experience in the future, and will more than likely ignore similar looking frogs. This is thought to be a "safety-in-numbers" mechanism that works by reducing losses due to curious predators that have yet to learn about poison frogs. For comparison photos, see Müllerian mimicry in Dendrobates frogs near Tarapoto, Peru

Agalychnis callidryas © Rob StegmannStartling Patterns - Shock Mechanism
Some amphibians, many of which are not actually toxic, possess certain physiological adaptations for the Rana warszewitschii © Carlos Cesar Martinez Riverapurpose of shocking potential predators, giving the frog a moment to leap away. This is most noted in the surprisingly large, red eyes of the frog Agalychnis callidryas, the red-eyed tree frog. During the day, this little frog camouflages itself by clinging to the undersides of green leaves, with any colored part of the body hidden from view. If disturbed, A. callidryas will suddenly open the huge, red eyes, which may be startling to the harasser, giving the frog a moment to escape. A. callidryas also possess bright colors about the sides of the abdomen, that could be passed off as aposematic coloration to unwitting predators. Bright colors are found on many non-toxic frogs, and are flashed at harassers in an attempt to confuse or fool them into thinking the frog is actually toxic.  The photo on the left shows A. callidryas with eyes open, and bright coloration showing. When sleeping, this frog closes a mesh-like lid over the bright colored eyes, and hides the orange feet and colored abdomen under the body.


Caecilian (Gymnophiona) Defense Methods


 
Toxicity
There is little known about the chemical properties of caecilian skin secretions, but studies show that their skin does produce biologically active substances. In Typhlonectes compressicauda (Typhlonectidae), toxins have been documented. This is an aquatic species whose predator, Hoplias malabaricus (Wolf Fish), is subjected to poisoning from T. compressicauda. Toxins have also been discovered in the skin of Siphonops annulatus (Caeciliidae). 

Tarichatoxin, Tetrodotoxin, and Salamanders


The poison found in in the granular glands of newts of the family Salamandridae is especially toxic, and was given the name tarichatoxin* after its isolation in western newts of the genus Taricha. Tarichatoxin* is biochemically identical to the water soluble alkaloid tetrodotoxin (anhydrotetrodotoxin 4-epitetrodotoxin, tetrodonic acid), or TTX, the third most potent non-protein neurotoxin known to exist (surpassed only by palytoxin and maitotoxin, which are approximately 100 times more poisonous than TTX). TTX is also very similar to saxitoxin, which is present in dinoflagellates (red tide) and responsible for shell-fish poisoning, but differs in its method of inhibiting the sodium channel.

Upon entering the blood stream, TTX will block the sodium channels of excitable membranes, causing paralysis in the nerves and muscles, during which the victim is usually fully conscience (Fuhrman, 1986; Yasumoto et al., 1986). Because of this unique characteristic, TTX containing animals have long been used by voodoo practitioners and others as a magic substance to turn people into 'zombies'. Toxic animal parts are typically crushed into a powdered and slipped into the victims meal or drink. One such account describes a Haitian man that was slipped this magic powder, and was pronounced dead as a result of his paralysis and inability to speak. When the TTX wore off several days later, the man tried to return home to his family, but was rejected because he was thought to be a walking-dead zombie.

Tetrodotoxin is found in poisonous Japanese pufferfish (fugu, of the genus Fugu), and in several other poisonous animals including globe fish, sun fish, trigger fish, the deadly blue-ringed octopus, frogs of the genus Atelopus, seastars, xantid crabs, horseshoe crabs, numerous marine snails, flatworms, and sea squirts, with more species still being discovered (Yotsu, et al., 1987; Yasumoto, et al., 1986; Noguchi, et al., 1987; Thuesen and Kogura, 1989). TTX has been found in four classes spanning four phyla, bringing up the question of where and how TTX is acquired by such diverse animals.

In 1984, Y. Shimizu's study of Japanese pufferfish showed that captive bred fugu lacked TTX, but acquired the toxin after consuming the livers of other TTX-possessing pufferfish. This led to the conclusion that the toxin was acquired from the natural habitat, i.e. the food chain (Yasumoto, et al. 1986). Shimizu hypothesized that TTX was acquired directly from ingestion of bacteria possessing the toxin, resulting in the toxification of the host organism in a symbiotic relationship (Shimizu, 1986). This would account for the wide variety of animals and habitats that TTX is found in. However, further studies by Yasumoto found that sympatric pufferfish similar to fugu accumulated TTX when ingested, but dissimilar sympatric fish did not. This finding seemed to contradict Shimizu's hypothesis. To add to the mystery, in 1995 Matsumura conducted a study on Shimizu's conclusions that dimmed the light on TTX's origins even more. Using technology that was unavailable in 1984, Matsumura discovered that captive bred fugu do in fact possess TTX, but in lower concentrations and in different tissues than wild fugu. The toxin in captive bred specimens is found in the skin and muscle tissue, whereas wild type fugu possessed high concentrations in the liver and gonads (Matsumura, 1996). Speculation as to the cause of such wide distribution and the origin of TTX continues today.

Although Matsumura's discovery seemed to contradict Shimizu's hypothesis, it is thought today that the toxin is acquired through the food chain from TTX-synthesizing bacteria. Through natural selection, pufferfish and other animals possessing TTX took advantage of a single-point mutation in their sodium channel that rendered them immune from the toxin. This enabled them to consume and adapt the poison to their systems without experiencing any ill repercussions. Similar spontaneous mutations are common in nature, but are not always advantageous to the organism.

Alteromonas spp. bacteria have been linked to the production of TTX in pufferfish, and several other marine species (Yotsu, et al., 1987; Yasumoto, et al., 1986). TTX has also been found in Vibrio spp. bacteria that were isolated from the intestines of xantid crads, tetraodontid fishes, and four chaetognath species (Noguchi, et al., 1987; Thuesen and Kogura, 1989). Alteromonas spp. and Vibrio spp. are thought to enter marine host organisms through the consumption of red calcareous algae (Jania spp.) possessing the bacteria. Vibrio spp. is thought to be the bacteria found in certain toxin amphibians, however, it is unknown how the bacteria may have reached them.

Although tarichatoxin is only 1% as potent as TTX, toxic newts are capable of inflicting serious damage, and even death, to large mammals and other potential wild predators. Interestingly, the common garter snake, Thamnophis sirtalis, of North America is immune to the toxic rough-skinned newt, Taricha granulosa, which it often preys upon. The two species seem to be in an arms race of sorts, for as the newts become more toxic to counteract the snake’s immunity, the snakes become more immune to counteract the increasingly poisonous newts.

It has been shown that all poisonous newts possess tarichtoxin, with Taricha species being the most toxic and Triturus species being the least. It has also been shown that terrestrial juveniles (efts) are several times more toxic than adults.

Batrachotoxin (C31H42N2O6), and other Dendrobatid Alkaloids


Batrachotoxin is found in many of the poison frogs of South and Central America. Like tarichatoxin, batrachotoxins are powerful, non-protein toxins, however, batrachotoxins are much more potent than tarichatoxin. A .00001 gram dose of batrachotoxin is enough to kill an adult human. Batrachotoxins effect the permeability of selective ions, resulting in depolarization of the nerves and muscles, arrhythmias, ventricular fibrillation, and possible cardiac arrest.

Batrachotoxin has long been used by South American natives for defense. By rubbing arrows and spear tips in the toxin secretions of poison frogs, namely Phyllobates terribilis, the Emberá and Noanamá Chocó Indians of Colombia have devised a powerful weapon, hence the common names "Poison Dart Frog", and "Poison Arrow Frog" commonly used to describe Dendrobatids. 

Frogs of the genus Phyllobates are the most toxin, with Phyllobates terribilis at the top of the list.

In addition to batrachotoxin, some species can produce other, less powerful agents, such as pumiliotoxins, histrionicotoxins, and gephyrotoxins. Pumiliotoxins occur in all species of the genera Dendrobates, and Phyllobates. Over 100 pumilio toxins have been isolated from both genera, which are divided into three groups: Pumiliotoxins A, B, and C. Pumiliotoxins A and B are considerably more potent than Pumiliotoxin C.

Atelopus frogs produce different toxins, for example, atelopidtoxin.


* - Tarichatoxin is essentially a dilute form of Tetrodotoxin (TTX), at around 1% concentration.

References: Journals

Catterall, W.A. 1980. Neurotoxins that act on voltage-sensitive sodium channels in excitable membranes. Ann. Rev. Pharmacol. Toxicol 20: 15-43.

Daly C.W., McNeal E.T., Overman L.E., et al. 1985. A new class of cardiotonic agents: structure activity correlations for natural and synthetic analogues of the alkaloid pumiliotoxin B (8-hydroxy-8-methyl-6-alkylidene-1-azabicyclo[4,3,0] nonanes). J. Med. Chem. 4:482-6.

Daly C.W., McNeal E.T., Gusovsky F., Ito F., et al. 1988. Pumiliotoxin alkaloids: relationship of cardiotonic activity to sodium channel activity and phosphatidylinositol turnover. J. Med. Chem. 2:477-80.

Garcia, Fuhrman, F.A. 1986. "Tetradotoxin, tarichatoxin, and chiriquitoxin: Historical perspectives." In C.Y. Kao and S.R. Levinson, eds., Tetradotoxin, Saxitoxin, and the Molecular Biology of the Sodium Channel. N.Y. Academy of Science 479: 1-14.

Garcia-Paris, Mario, and Stephen M. Deban. 1995. A Novel Antipredator Mechanism in Salamanders: Rolling Escape in Hydromantes platycephalusJournal of Herpetology 29(1): 149-151.

Noguchi, T., D.F. Hwang, O. Arakawa, H. Sugita, et al., 1987. Vibrio alginolyticus, a tetradotoxin-producing bacterium in the intestine of the fish Fugu vermicularis vermicularis. Marine Biology 94: 625-630.

Thuesen, E.V., and K. Kogura, 1989. Bacterial production of tetradotoxin in four species of Chaetognatha. Biol. Bulletin, 176: 191-194.

Yosumoto, T., H. Nagai, D. Yasumura, T. Michishita, et al., 1986. "Interspecies distribution and possible origin of tetradotoxin." In C.Y. Kao and S.R. Levinson, eds., Tetradotoxin, Saxitoxin, and the Molecular Biology of the Sodium Channel. N.Y. Academy of Science 479: 1-14.

Yotsu, M., T. Yamazaki, Y. Meguro, A. Endo, et al., 1987. Production of tetradotoxin and its derivatives by Pseudomonas sp. isolated from the skin of pufferfish. Toxicon 25: 225-228.

References: Books

Hofrichter, Robert. Amphibians: The World of Frogs, Toads, Salamanders and Newts.
   Firefly Books, 2000.

References: Printed Articles

Light, Willian Haugan. 1998. Eye of Newt, Skin of Toad, Bile of Pufferfish.
   California Wild, Summer 1998 ( California Academy of Science).


Sheppard, Jessica. 1999. California's Noble Newt.
   California Wild, Winter 1999 ( California Academy of Sciences).

References: Internet

Yoshida, Kari. Anti-Predator Mechanisms in Salamanders. University of Oregon. http://www.uoregon.edu/~titus/herp/antipredator.htm (Accessed: 2002).

 

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