The Texan Herper: Amphibians

AMPHIBIANS

DISCUSSION:
The word 'amphibian' refers to living two lives in water and on land. All amphibians are non-amniotic vertebrate animals, which means besides having no yolk, the egg is not terrestrially adapted. Therefore, eggs are laid in water or at least moist environments and in also being non-amniotic, there is no limb development during the fetal stage. This in turn necessitates a larval stage upon hatching. Larvae must go through metamorphosis to attain adult form.

1. The extinct Golden Coqui

There are, as always in most cases, exceptions to the rule. Toad species belonging to the genus Rhinella produce amniotic eggs on land albeit in humid environments. The toadlets hatch directly from the egg foregoing the tadpole or metamorphose stage. All treefrog species belonging to the rather large genus Eleutherodactylus lay eggs on palm leaves or on other moist terrestrial plants and guard the eggs. In some cases, once the developing embryos are hatched, the adults (in some species the female and in others the male) for a short duration will also guard the little froglets which possess minute tails that shortly are absorbed into the body. In fact, one species in this genus, the recently extinct Golden coqui (Eleutherodactylus jasperi) gave birth to live young and exhibited parental care by attending to the newly born froglets. Freeing them from being bound by water restrictions, nature has allowed these toads and frogs to live in higher altitudes and in tropical jungles further from access to permanent water sources.

Further exceptions are the plethodontid salamanders that lay terrestrial eggs with fully developed but miniature hatchlings, along with caecilian species that give live births to viviparous or ovoviviparous offspring.

The amphibious line first made its appearance during the tail end of the Devonian period around 375 million years ago (mya) and became the ruling predator into the Carboniferous and Permian Periods; a span of around 100 million years. The earliest amphibian in the fossil record to date is the late Devonian, Elginerpeton found in 368 million year old rock near Scat Craig, Scotland. The fairly well known Ichthyostega was discovered in 363 million year old Greenland deposits.

2. Elginerpeton

Though these two animals were definitely true tetrapods with amphibious features, they at best could be considered very primitive amphibians. In other words, the Amphibia class was trending but was not wholly yet a true amphibian.

3. Ichthyostega

The late Devonian was a period that witnessed a great greening of land with multitudes of plant species diversifying out and away from permanent water sources with root systems aiding in the further erosion of rock into soils. Primitive arthropods co-evolving with the diversified vegetation, evolved into the first true insects that shared in the mutual evolvement and dispersal of seed bearing plants; much like pollinating insects and pollen bearing plants do today. Honey bees today, pollinate over 30% of our edible plants.

This great plant life takeover of terrain created a carbon dioxide sinkhole. In taking the gas out of our atmosphere, it produced a cooling climatic trend that perhaps was one of the main causes of the massive extinction event witnessed during the late Devonian, primarily in the marine communities and in particular, species in shallower waters such as corals. One group of shallow water animals were not affected by the extinction and may have benefited from it. These were the lobbed finned fishes.

Lobe-finned fishes differ from today's more prevalent ray-finned fish in that instead of fins, have two paired fleshy lobes supported by internal phalange bones. Devonian lobe-fins also had developed skull spiracles to gulp in air possibly to survive oxygen poor shallows or a duration of droughts.

Some of these fish, like the extinct Panderichthys of northern hemispheric oceans and Gogonasus of southern hemispheric oceans, would venture out onto shorelines walking on their lobes. Being a truer fish in anatomy than in being a terrestrial tetrapod, they were greatly dependent on a watery environment. Nonetheless, even though it was on lobed-fins rather than true legs, they were the first vertebrate to walk on land. Evidence of a copious and vast amount of untapped food resources in terrestrial plants and insects perhaps induced evolving adaptations to become more terrestrial and in between 400-350 mya, the evolution of true tetrapods evolved from the lobed fins of these fish.

375 mya, Tiktaalik appeared and though not a true tetrapod, was a transitional prototype tetrapod that wasn't fully a fish either. Though classified under the Sarcopterygii class of fish, this animal had possessed weight bearing wrists that could flex, simple ray bones reminiscent of fingers and the first fish ever to lack bony plates in the gill area, thus restriction of lateral head movement was eliminated. This essentially gave the animal a neck with the pectoral girdle separate from the skull giving more latitude in hunting prey on land.

In addition, Tiktaalik had a supportive rib cage applicable enough for body support over land and tetrapodal lungs. Eventually Tiktaalik and other prototype tetrapods gave rise to the first true tetrapods...the labyrinthodont. Labyrinthodonts like Ichthyostega and Acanthostega, 365 mya developed limbs with digits on the four hands. Though Acanthostega possessed scales on the gills, both species had skin coverings instead of scales that would protect them from land desiccation. In addition, their mouth and teeth were adapted for terrestrial style feeding. Still though, they spent more time in water than not.

Finally, around 310 mya, amphibians were firmly established, with dominant examples showing up in the fossil record, as in the appearance of the species Eogyrinius, which is Greek for dawn tadpole. This creature was huge for an amphibian growing to 4.55m/15ft. With stumpy but fully developed legs, the amphibian was semi-aquatic and appears to have prowled shallow river beds much like crocodilians do today.

4. Eogyrinus

5. Eryops

One of the best known of prehistoric amphibians is Eryops who showed up in the Permian around 295 mya. Eryops, with massive limbs and a short tail was most probably by far more terrestrial than aquatic. This 2m/6ft amphibian had true ligament attachments in the binding of the ilio-sacral structure and though it may not have been phylogenetically linked, it no less represented an intermediate stage between the primitive amphibians and early reptiles. This ligamentous attachment within the limbs of Eryops was a definitive stage as the precursor to cartilaginous variations utilized in all modern terrestrial animals for limb locomotion.

6. Skeletal Amphibamus

7. Fossilized Amphibamus

8. Amphibimus

In all actuality, the true final juncture of modern day amphibians before they had split up into frogs/toads and salamanders/newts is found in the much smaller amphibian, Amphibamus, which was no larger than 15.2cm/6in. Amphibamus lived during the late Carboniferous around 300 mya.

Modern amphibians are technically classified as lissamphibians comprising frogs, toads, salamanders, newts and caecilians. Lissamphibian radiation appears to go back to the insectivorous Amphibamus as the common ancestor. Lissamphibia is now a technical subclass for today's amphibians.

Amphibian main traits are four limbs with digits ending in toes or webs with no claws or nails, lungs taking over gill function, possess internal as well as external nares (nostrils), smooth moist skin made to absorb dissolved oxygen, keratin (a protein) in skin to counter dehydration, ectothermic, exhibit torpor through hibernation, aestivation or brumation and have an aquatic larval stage that metamorphoses into the adult stage.

Metamorphism is controlled by the produced hormone, thyroxine. What is interesting here, is that there are certain amphibians, such as the axolotl that remain in their larval stage throughout life. This state is termed, neoteny and even humans possess some forms of neotenic stages. But if iodine is introduced into the larva of a neotenic salamander, it will then metamorphose and fully develop into the adult form. Thyroxine is dependent on iodine where low levels of the element thwart growth. But with the introduction of iodine, the higher levels promote the metamorphosing stages.

Apodan Characteristics (Caecilians)

Tropical-semitropical burrowing worm-like animals.
Small or vestigial eyes, most are blind.
Completely lack external limbs
Only amphibian to internally fertilize
Most species bear live young

Caudata Characterisitcs (Salamanders/Newts):

Smooth moist skin for cutaneous respiration.
Elongated body with a tail and normally 4 legs in adults.
More water dependent than anurans overall.
Nocturnal in drier regions.
Contains completely aquatic species such as newts, amphiumas and sirens
More species bypass metamorphism stages than anurans with some species giving birth to live young.
Eggs usually require water for development from submersion to dampness.

Anuran Characterisitics (Frogs/Toads)

All adults without tails to facilitate jumping or hopping.
Most go through metamorphism.
Typically frogs have smooth skins and more dependent on water while toads have warty dry skin.
All anuran tadpoles have gills and tails.
Long hind limbs for frog jumping and toad hopping.
Possess a long forked tongue hinged at the front of the mouth.

Classification:
The classification outline here for amphibians is going to be a different format than what was approached in reptiles. In so doing, I'm attempting to expose the various styles of nomenclature and with the introductions, you can decide on which form of classification suits you best. There are many out there in varying outlines and systems.

NOTE:
In this classification scheme, on the very top I have included the order, Trachystomata (also known as Meantes) for sirens, because the trending is to list the family, Sirenidae in a different order as separated from salamanders altogether. The reason being is that through new genetic analysis, sirens are not phylogenetically related to any other salamander group. Many of their unique primitive and derivative characteristics are from a distinct line. So, I have listed Trachystomata/Meantes as the distinct order of sirens, but still included the old version under Caudata grouped with the salamanders. Still further for some form of distinction, under Caudata I added the suborder, Sirenoidea to show a separation of sirens from the rest of the salamander groups listed under Caudata.

The discussion above brings us to Caudata. Caudata was the original salamander order, but now it is becoming common to list all extinct salamander forms under Caudata and all extant species under the new order, Urodela.

There is as well another item. Originally mudpuppies, waterdogs and olms were all listed under the one family, Proteidae simply because they all are neotenic and remain in water with gills their entire lives. But due to their unique adaptations and features from living submersed as troglobites (animals who spend their entire lives in caves), olms remained under the family, Proteidae while mudpuppies and waterdogs were removed to the new family listing of Necturidae that coincides with their genus, Necturus.

I hope that this does not create confusion. Science is flexible and though old ways sometimes die hard, science will change when the change is warranted. It's just that during the transition, the water can get a little murky, if not downright muddy.

There are two main amphibian classification schemes today and they are Frost: Amphibian Species of the World (ASW) and AmphibiaWeb by the University of California (Berkeley). The two schemes differ greatly in number of families.

Where AmphibiaWeb stands more traditional, Frost ASW has split several families off from original families and elevated them to distinctive families.

Under the class, Amphibia as listed by Frost ASW, there are 3 orders with 61 families.

Following and covering some 6,775 species is the taxonomic AmphibiaWeb scheme in amphibian classification:

Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Class: Amphibia

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Sirens:

As mentioned earlier, certain taxonomic classifications for sirens are under the order, Caudata catalogued with the rest of the salamander groups, but with the additional stipulated suborder, Sirenoidea. Due to unique phylogenetic characteristics, reclassification is warranted and the current trend is to list them under a separate order known as Trachystomata, but sometimes listed as Meantes. Try and keep this in mind when reviewing other amphibian classifications.

Sirens make-up only one family, Sirenidae with two genera. The genus, Siren consists of two species while the genus Pseudobranchus (dwarf sirens) has seven species listed. Though there is disagreement on the actual number, tentatively there are five Psedobranchus subspecies and three Siren subspecies. Sirens only occur in the southeast portion of the U.S. and a portion of northern Mexico.

All these animals have small but fully developed forelimbs while any hindlimb or appendage is totally absent in also lacking a pelvic girdle. Although they are listed as being neotenic, in a sense they show neotenous reversals in that larval gills, in contrast to usual amphibian pedomorphism, are very small and non-functional relying on lungs, whereas in adults the gills become fully formed and functional, though still retaining the developed lungs. This hints to a consensus that sirens originated from a terrestrial ancestor that possessed an aquatic larval stage. Developed larvae, may even shrink their gills to stumps and rely fully on lungs for breathing by gulping surface air if the body of water they're in is too stagnant and anoxic (the depletion of dissolved oxygen).

These animals range in size from the northern dwarf siren (Pseudobranchus striatus) averaging 10.2-21.6cm/4-8.5in in length to the greater siren (Siren lacertina) reaching lengths of 97cm/38in.

1. Northern dwarf siren ~ P. Striatus

Sirens live in marshy murky waters, bottom lands and even ditches that have tendencies to dry out especially during drought stages, but are also found in more permanent bodies of waters like large ponds and muddy slow moving streams. Though small, the two front legs are capable to move the animal across land during rainy nights to reach wetter habitat.

If no water source is reachable during droughts, sirens will burrow into drying mud, encase themselves into a moist retaining mucus cocoon and lower their metabolism and heart rate drastically. The cocoon is leak proof and keeps them from desiccation. Once cocooned, they switch from gill breathing to lungs and wait through aestivation for the next rains to come and fill their habitat back up with water again.

Even though they can extremely lower metabolism down to 30% functionality, metabolic rates must be high enough during active periods, for unique among amphibians, they can tolerate much colder conditions. They are commonly observed swimming just beneath ice during winter months.

2. GB heron & greater siren

The only teeth present are small and few located on the palate and splenial bone positioned on the lower jaw. For biting purposes, there is no dentition, but instead a horny sheath much like a beak is utilized in defense and for capturing prey. Diets mainly consist of invertebrates and filamentous alga; another unique siren characteristic in an amphibian's diet consisting of vegetation. Some filter feed in the detritus mud bottoms, while others will hunt and prey on terrestrial invertebrates during warm rains.

Natural predation has not been fully studied, but wading birds, water snakes and adult fish have been observed preying on sirens. Eggs and larvae are preyed on by fish, frogs and even sirens foraging on water bottoms consume siren eggs.

Two other unique features of the body that distinguishes sirens from the rest of the salamander groups is that sirens possess an interventricular septum in the heart common in mammalian hearts and at least two species can produce vocalizations. Most other sirens, though they don't have vocalization, do communicate with each other by using a series of clicks for intraspecific conveyance. The clicking process is performed by gulping surface air, then jerking the head up and down sharply. The forced expulsion of air and head movement causes horny coverings in the mouth to clip together creating the clicking. Rates of clicking are controlled.

Sirens are the most ancient line of extant salamanders and are listed together with the other living salamanders under the order, Urodela. In this taxonomic scheme all extinct salamander forms are listed under the order, Caudata. Sirens are further divided out in only one family, the Sirenidae with two genera and four species.

1. Southern dwarf siren ~ P. axanthus

Southern Dwarf Siren
Family: Sirenidae
Genus: Pseudobranchus
Species: Pseodobranchus axanthus

Length: 10-25cm/4-9.8in
Weight: ~ 69.45g/2.45oz

Longevity: Under ideal conditions, captives have lived 25 yrs; in the wilds would be considerably less.

Name Origin: Pseudobranchus axanthus ~ is Greek, 'pseudos'-without, 'xanthos'-yellow; meaning without yellow in color comparison to other specie of dwarf siren, P. striatus.

Distribution: Is endemic to the southeastern half of Florida found in wetlands shallow sinkholes and cypress ponds containing dense mats of water hyacinths.

Description: Are a thin unusually slimy salamander, even for salamanders. Have both lungs and gills. The prevalent gills are feathered and bushy, often visually hiding the tiny forelimbs that end in three toes. No hind limbs. Coloration varies but are generally brown and sometimes black or gray, but always with three dull yellow or tan longitudinal stripes running down the entire length of the back and sides. There was a trend to link the two species and all subspecies into one, but molecular genetics proved they were indeed distinct animals having different chromosome counts; there are 2 subspecies.

2. P. axanthus displaying gills

Diet: Has a small blunt rounded mouth so eats any invertebrate prey small enough to take in, including worms, ostracods, chironomids and amphipods.

Habits: Is nocturnal preferring to stay hidden under debris or mud on shallow water bottoms; larvae will often congregate around stem roots of hyacinths; little is known about reproduction; spawning has been observed from November to March with eggs placed and attached singly to aquatic plants; eggs have 3 jelly layers; male captives have been seen to follow the laying female then wrap around an egg once she has deposited it; suspect this is the way he inseminates, but no free swimming sperm is ever noted; produce faint yelps when caught.

1. Siren intermedia texana

Rio Grande Lesser Siren
Family: Sirenidae
Genus: Siren
Species: Siren intermedia
Subspecies: Siren intermedia texana



Length: 69cm/27.2in (largest of any siren subspecies)
Weight: 312g/11oz
Longevity: 4-7 yrs

Name Origin: Siren intermedia texana ~ is Latin referring to the 'aquatic sirens' of Greek mythology that sang alluding to salamander siren's possessing a voice and 'intermedia' denoting 'intermediate' between the greater and dwarf sirens in size; 'texana' simply refers to 'of Texas.'

Distribution: Found within the lower Rio Grande River watershed in still and slow moving water bodies that are usually weed choked and murky. Preferring any shallow body of water keeps them from fish predation found in deeper clearer more permanent waters. Extensive usage of pesticides, herbicides and other pollutants are contaminating the preferred stagnant water bodies threatening this subspecies; there are 3 subspecies of intermedia.

Description: The dorsal, dorsum or top side, whatever is your choice in calling it, is a dull gray to brownish dark gray; ventral, venter or belly side is an even light gray; has the most costal grooves of lesser sirens exhibiting up to 38; body plan is very eel-like in appearance with two small front legs each with four toes; small, but useful beady eyes.

Diet: Will take about any invertebrate it can grab or suck in by or with the mouth. Eats terrestrial or aquatic insects, snails, worms, crustaceans and an occasional small fish if available. Quantities of aquatic vegetation material have been found in dissected stomachs, but is apparently not easily digested.

Habits: This siren is passive by day preferring to lay just buried in mud or sand bottoms, or burrowed in plant debris that has settled to the bottom; is primarily nocturnal and at night is most active in foraging or pursuit of other sirens; along in breathing with gills will also come to the surface to gulp air; bite marks from other individuals are common on both sexes, so courtship must be a bit violent; female lays upwards to 300 eggs in the water; larvae are small at 1.1cm/.04in and take upwards to 4 years to mature; give off a clicking sound when in company with other sirens and may utter a shrill cry when seized by a predator or a human's hand.

Vulnerable 3.1(IUCN)

1. Greater siren ~ S. lacertina

Greater Siren
Family: Sirenidae
Genus: Siren
Species: Siren lacertina

Length: 48-97cm/19-38in
Weight: Are the heaviest N. Hemispheric salamanders weighing over 1kg/2.20lbs
Longevity: Life expectancy in wilds is not known, but captives have lived 25 yrs
Name Origin: Siren lacertina ~ Latin, 'siren' pertaining to Greek temptress mythology that sang and 'lacertina' for serpentine body.'

Distribution: From southern Maryland hugging the eastern coast seaboard down to Florida

2. S. lacertina showing lighter ventrum

Description: Is the most heaviest and third largest salamander in North America; coloration is an overall drab gray or olive with random dark spots on head, sides and back; younger greater sirens have a light stripw down each side that fades with age; the ventrum and sides are lighter with faint greenish yellow shows; eel-like in appearance; has a rudder shaped tail with two fairly small front limbs possessing 4 toes; no hind limbs; small beady weak appearing eyes; is the most primitive of extant salamanders; there are no subspecies.

Diet: Being the largest of sirens the diet may be accommodated more with larger food; in addition to aquatic and terrestrial invertebrates and their larvae, will also consume tadpoles, other salamander larva, aquatic eggs, small fish, crawdads (crayfish) and on occasion plant matter; utilize a lateral line (a sensory organ running laterally down each side of the body that detects vibrations and movement) even as adults to detect prey; most amphibian larvae possess a lateral line as inherited from fish lineage, but lose it during metamorphosis; stores fat in tail and along vertebral column for dry seasonal nourishment.

Habits: Is nocturnal, spending days submerged and burrowed in debris or mud in bodies of shallow water; will come out on land during rainy nights; are sexually mature on average at two and a half years of age; females will lay from 100 to 500 eggs singly or in small clusters attached to leaf litter submersed in shallow waters; with the ability to aestivate this siren can lose up to 85% of its body weight and still survive; can tolerate brackish waters; aquatic herbicides and man induced drainage of wetlands are making sirens less common throughout its range, though in localized protected areas may be common.

Caecilians:

Caecilians, mistakenly known colloquially as 'blind snakes' are not just an odd amphibian, they are simply just one odd critter period. Being totally legless and long, the smaller species look like earthworms, even displaying the skin fold body rings (annuli) of segmented annelids, while the larger ones resemble snakes. Sometimes they're even called the 'two-headed snake', because their blunt tail-end (in most cases, a true tail is absent) is not well defined. Eyes on the head are either very tiny or hidden altogether under flesh and bone. Fossorial or aquatic in nature, evolution has left them with degenerative eye muscles and optic nerves. The retina and crystallized eye lens are vestigial at best and in the scolecomorphids of Africa, the entire eye has been pulled out of socket and predisposed under skin of the upper jaw. So yes, for the most part they are blind to vision, but do distinguish light from dark.

Caecilians may appear as slow and sluggish, but they can react with lightning fast speed to a stimuli and can travel about as quickly as any snake can.

In rearing young, skin feeding is carried out by the mother in members of the genus, Boulengerula and Siphonops. Known as maternal dermatophagy, this is where the caecilian offspring actually tear into the mother's flesh, rip off chunks and eat it. It sounds gruesome, but it does save the expense in spent energy of maternal care in hunting, capturing and bringing back prey to be consumed by the offspring. Also, during this period, it appears that Mother Nature is not just looking out for the offsprings' welfare but for mom's too.

For the offspring's sake, Mother Nature has evolved special 'infant teeth' specifically designed for ripping into the mother's flesh. These teeth are later lost and replaced as the caecilian matures. Some of these infant teeth are spoon shaped for scraping while the rest are spiked points for piercing outer skin layers.

For poor mom's sake, Mother Nature has equipped her to withstand the onslaught. During this period of rearing young, the mother's skin doubles in thickness with several layers. Only the outside layer is eaten and appears to not be equipped with nerves to signal pain.When the outermost layer is scraped and ripped off, the next layer quickly replaces it, maturing into the next meal. The mother does lose weight, but otherwise appears unharmed during this weaning stage.

What is truly remarkable is that caecilians species are oviparous (egg layers), ovoviviparous (eggs hatch inside female, then born live) and viviparous (give live birth) and it doesn't matter, all birth types, except for oviparous species with aquatic larval stages, participate in some form of maternal after hatching or birth care. In addition though, in live bearing caecilian forms, the offspring, while still inside the mother will scrape off nutritious tissue and fatty secretions from the linings of the female's reproductive organs with the infant teeth.

This has to be the most purest form of parental investment but is not so far out there in extremes. It does parallel mammalian strategies in rearing offspring, for a big investment in mammalian mothers in producing and offering nutrients from her own body it seems, has similar infant rearing strategies. In a sense, whether it's skin from the body of a caecilian parent or milk from the body of a mammalian parent...it is an invested strategy in the successful rearing of a totally dependent offspring.

With caecilian habitat spread out across the globe, they hail from the the wet tropical regions of east and west Africa, Southeast Asia, India, the island Sri Lanka, the Seychelle Islands, in portions of northern and eastern South America, in parts of Central America and subtropical Mexico. They are only found in moist environments.

Except for one family being aquatic, caecilians are strictly fossorial, living a subterranean existence throughout their lifespan burrowing in moist soils in search of worms, termites and other small invertebrates for diet concerns and for each other in mating and rearing of young. The whole body plan is designed to be fossorial.

Expounding a bit more on what was mentioned earlier in comparing them to earthworm body plans, caecilians have numerous ring-shaped folds in the skin known as annuli that partially encircle the body. This gives the appearance of segmentation and allows for a more proficient propulsion in tunneling.

Caecilian bodies are covered in a mucilage slime. The mouth is recessed and has two, not one as is the case in most animals, jaw muscles that keeps the jaw and skull rigid. These arrangements are to aid in keeping soil from entering the mouth. In species with eyes, skin covers them for protection while external ears are absent. One other aspect in burrowing efficiency is that caecilians do have calcite scales, but are internal, covered by a smooth layer of external skin.

Hearing perception is restricted to picking up and interpreting ground vibrations by means of an auditory ossicle (small hearing bone) located in the skull and articulated with the jaw. Collectively, this apparatus arrangement is known as the collumella.

The skull is bullet-shaped and sturdily fused for proficiency in tunneling. Except for eye, nare and tentacle openings, the skull is completely roofed and is a condition known as stegokrotaphy. A few species however exhibit zygokrotaphy; a condition in having an opening in the temporal region facilitating kinetic movement of the skull.

1. Caecilian (Dermophis mexicanus) skull & teeth

For olfactory reception, most caecilians have nares (nostrils) for picking up airborne particles and in addition, have a pair of tentacles connected to optic nerves and muscle that are set along grooves between the eyes and nares and are extendable and retractable for detecting molecules left by prey passing through the soil. Borrowed from the fish lineage, caecilian genera with an embryo or larval stage possess lateral line organ systems embedded in the skin on the sides of the body and head for mechano-reception (neuromasts) and electro-reception (ampullary organs). Adult aquatic and all viviparous caecilians, lose this trait.

Caecilians are the amphibian with true teeth in having recurved sharp teeth as snakes do and use them for grabbing onto prey, such as worms, termites and grub worms (beetle larvae). Also alike as snakes, there are two rows of upper teeth while the lower teeth line the jaw and fit in place between the upper rows when the mouth is closed. Teeth are strictly used for grasping and not chewing.

Viviparous caecilians have fetuses that possess fetal teeth. It was first thought viviparous caecilians came about strictly from convergent evolution, but possessing fetal teeth suggests otherwise and implies that viviparous caecilians evolved from oviviparous ancestors that already had specialized in dentition which was preadapted to feeding in oviducts.

Also, as in snakes, most caecilians breathe with lungs with the left lung atrophying while the right one is expanding and is highly compartmentalized. Caecilians with lungs take in air through the mouth and skin to absorb oxygen in the lungs. Cutaneous gas exchange (skin breathing) is possible due to a permeable thin layer of skin intermixed with dense capillary networks that can absorb oxygen and emit carbon dioxide. Some caecilians for lungs, have nothing more than an extended trachea.

There are two species of caecilians that have done away with lungs completely. One is aquatic and one fossorial with both utilizing skin respiration, but in addition, Atretochoana, the aquatic genus has had its gills fused into a sac-like structure. Both lungless species have also lost the use of their nares in being internally sealed. The photo below depicts the newly discovered lungless terrestrial species, Caecilita iwokramae.

2. C. iwokramae ~ discovered in 2009

A tail is any part of the body going beyond the cloaca, but a true tail is supported by vertebrae. With this definition in mind, there are only two caecilian genera then that would have a true tail.

Male caecilians have a penis-like appendage known as the phallodeum, in which he inserts into the female's cloaca and inseminates her internally after an average of 2 1/2 hours of copulation.

In most, there is no need for metamorphosis, for the young are hatched or born fully formed. In one genus only, there is a short aquatic larval stage and still, there are a few species with larvae that are not aquatic spending time instead in very moist soils. Some young are hatched or born with gills, but quickly lose them.

Maternal care is obvious throughout caecilian families with oviparous females incubating and protecting her clutch of large yolk filled eggs by wrapping around them. Some egg laying caecilians will further perform post-paritive parental care for the young including the practice of maternal dermatophagy, where the young (pups) feed off the mother's outer skin layers. Ovoviviparous females have the eggs hatch still within her where the young remain until they mature. As in the case of oviparous hatchings, ovoviviparous young are also equipped with infant teeth and rasp off and eat oviduct cells from the mother while maturing inside her.

3. Eocaecilia micropodia

The fossil record is difficult to find for these burrowing creatures, but the extinct caecilian genus, Eocaecilia that lived during the early Jurassic shared characteristics with true salamanders and the extinct burrowing species of the order, Microsauria. Unlike modern-day caecilians, Eocaecilia possessed small legs and developed eyes showing extant caecilians originated from a terrestrial lineage.

One thing that baffles me and I'm sure quite a lot of herpetologists and that is why do a lot of caecilians have color? Though the general color is a drab gray, many species are vibrantly colored or varicolored. For an animal that spends most its time in total darkness underground and is for the most part essentially blind, why then the expense in wearing a colored skin?

To take a gander in guessing, I suppose it could be the result of aposematism. Aposematic animals are those that are toxic and display color warnings in advertising to predators not to eat them unless wanting to pay for it later. All caecilians possess toxic granular (poison) glands embedded in their skin that secretes poison onto the outer epidermal layers. Perhaps the more colorful ones carry more potent toxins and inherited the color from their once terrestrial ancestors. The toxin also protects oviparous caecilian eggs acting as a disinfectant to bacteria and fungi.

Six families, thirty-six genera and ~ 171 species make-up the caecilian groups under the order Gymnophiona (Apoda). Family rankings are currently in a period of transition in making up new ones. Dermophiidae, Herpelidae, Indotyphlidae and Siphonopidae, as being borrowed from Caeciliidae are now being given their own ranking as paraphyletic to Caeciliidae, but, monopyhletic unto their own distinct morphology and genetics. Specie members of these new families are most probably a derivative of allopatric function as determined from the latest genetic data.

An exception to this monophyletic family trending are the two species in the genus, Herpele which are the only remaining evolutionary species from an ancient and extinct lineage. The two Herpele species are far removed from their closest genus relations, Boulengerula, even though they are now both being listed under the trending family, Herpelidae.

With this in mind the six standard caecilian families are: 1) the common caecilians (Caeciliidae); 2) the aquatic caecilians (Typhlonectidae); 3) the fish caecilians (Icththyophiidae); 4) the tropical caecilians (Scolecomorphidae); 5) the beaked caecilians (Rhinatrematidae) and 6) the Indian caecilians (Uraeotyphlidae).

This may all be a bit confusing, but until 1968, there was considered only one family of caecilian. Newer information from better and more updated technology has brought caecilian research and knowledge a long way with sufficient data in understanding these creatures physiologies, morphologies and attributes.

1. Atretochoana eiselti

Atretochoana Lungless Caecilian

Family: Caeciliidae
Genus: Atretochoana
Species: Atretochoana eiselti

Length: 738mm/29.05in (largest lungless tetrapod known)

Weight: Estimate ~ 130-200g/4.8-7.4lbs

Longevity: Unknown
Name Origin: Atretochoana eiselti ~ 'Atretochoana' is Greek for 'closed nostrils;' 'eiselti' after Josef Eiselt (Austrian herpetologist).

Distribution: Endemic to Brazil in higher elevation, fast flowing and clear smaller rivers and permanent streams.

Description:There are only two preserved specimens with no living population known; readily distinguishable form other caecilians by possessing wrinkled skin, massively large mouth for caecilians and internally sealed nares due to choanal apertures (passageway from back of nose to throat) being permanently sealed with fleshy flaps; head is broad and flat being dorsoventrally compressed; eyes seated in shallow ocular depression; high number of splenial (thin splint-like bone) teeth; lungless utilizing cutaneous gas exchange; in addition, has mobile cheeks and stapes from which muscles originate that is unknown for any other species; a well developed fleshy and short dorsal fin runs down the back; coloration is dorsally a light blue-gray while lighter ventrally with a white patch on ventral surface of head; tip of snout and mouth borders are olive-gray; there are no subspecies.

Diet: Unknown; only one of the two preserved specimens had stomach contents and that was small quartz crystals; assumed to eat small fish and aquatic invertebrates.

Habits: Other than being aquatic and possibly viviparous, nothing is known about its life history or behavior.

Data Deficient (3.1) IUCN

1. Sao Tome Caecilian ~ S. thomense

Sao Tome Caecilian
Family: Caeciliidae
Genus: Schistometopum
Species: Schistometopum thomense

Length: 135-345mm/5.32-13.58in; up to 375mm/14.76in.
Weight: Averaged 136g/4.80oz.
Longevity: Unknown; due to smaller size perhaps 5 plus yrs.

Name Origin: Schistometopum thomense ~ Schistometopum is from a Latin neuter noun meaning 'split forehead;' thomense is in refernce to geographic range.

Distribution: Found on the island of Sao Tome in the Gulf of Guinea 220km/136.7mi off the coast of West Africa; reportedly, once shown S. thomense photos local Ituri Province natives of the Democratic Republic of the Congo on the African mainland, said they recognized and had seen specimens of this caecilian; prefers softer soils and pre-existing burrows in steep terrain; uncommon in drier northwestern portion of island; in wet conditions is more apt to surface and can be found on ground, under superficially loose soil and under rotting vegetation floor litter; drier conditions drive them deeper into soil; infrequence to agricultural plots; based on genetic variance this caecilian had already inhabited the island prior to human settlement; is a mystery how S. thomense inhabited the island; Sao Tome is a shield volcano that arose from the ocean floor, but is a chain of volcanoes that perhaps the caecilian island hopped on debris rafts originating from the mainland.

2. Note dermal scalation

Description: Eyes visible through skin and are not in bone, but embedded in squamosal sockets; splenial teeth present; dermal scales present, in particular in anterior just beneath the skin; tentacular aperature closer to eyes than nares; lacks a true tail; head is sloped from crest of forehead to terminus of upper mandible; mouth is ventrally recessed; sexual dimorphism is not pronounced in total length, but in head size; the adult male head is larger than the females; at ~130mm/5.12in sexes begin to exhibit dimorphism in head shape with male's expressing blunter more larger heads than females; dorsolateral coloration of uniform bright yellow to darker burnt yellow; nonuniform and random whitish ventrum; purplish brown flecking may or may not occur; chromatic flecking is more common in morph specimens south of their range and those lacking the flecking are more common in the northern tier of their range; there are no subspecies.

Diet: Predominately preys on earthworms supplemented by centipedes, ants, insect larvae and mites; juveniles primarily prey on endogeic invertebrates (animals that range just below the ground surface), in particular small earthworms; rather than pursuing prey prefers the lie in wait ambush approach with a positioned burrow posture of head raised slightly above and perpendicular to burrow floor and neck cocked forward all the while remaining motionless; once seizes and subdues passing prey, utilizes long-axis rotation to tear off chunks of prey for ingestion; due to noxious taste from toxic secretion, only ants in numbers seem to be the primary predator.

4. S. thomense preying on earthworm

3. Adult with neonates

Habits: Are more surface or near surface dwelling than most caecilians; in soil, will excavate their own burrows and tunnels and keep, maintain and use them for monthly periods before new excavation commences; will also use passages in soil created from rotted out tree roots; below ground locomotion is achieved through a combination of internal and whole body concertina locomotion; above ground a combination of lateral undulation and internal concertina locomotion results in movement; have been observed moving with ease backwards through tunnels when necessary; are mainly nocturnal; primary breathing is cutaneous respiration through highly vascularized skin; additionally breath with pulmonary respiration; toxic skin gland secretions, whether from contact or fuming in a confined area, are potent as other animals contained in the same holding tank with this caecilian die from toxin's exposure; apnea has been observed up to 3 hours due to low metabolic rate at rest and highly efficient cutataneous gas exchange; bite marks and teeth scrapping scars are common on both adult sexes and juveniles; appears to be more the result of a chemoreceptor communication process tied to the vomeronasal activity of the tentacle than a mating ritual or territorial dominance duel; no stage in reproduction is tied to water; captive females reproduce biennially and are viviparous producing live births of 2-7 neonates (young); fetus possess an infant tooth and nurses through intraoviductal feeding; neonates are born ranging from 91-118mm/3.58-4.65in and double in size within nine months; neonate siblings will loosely knot together in enlarged tunnels.

1. Tapalcua ~ D. mexicanus

Tapalcua
Family: Caeciliidae (Dermophiidae)
Genus; Dermophis
Species: Dermophis mexicanus

Length: 30-50cm/12-20in
Weight: 109-179g/3.85-6.31oz
Longevity: 8-11 yrs, Maximum 13.8 yrs
Name Origin: Dermophis mexicanus ~ Dermophis: dermis is a latinized form of the Greek word derma for 'skin;' ophis refers to the Greek word, 'ophidian' meaning 'serpent;' mexicanus refers to a portion of the species' geographical range.

Distribution: Southeastern & southwestern Mexico down through the Central american countries of Guatemala, El Salvador, Honduras and Panama; found in lowlands and mountains of Guerrero, Mexico in the western Pacific versant and Veracruz, Mexico in the eastern Atlantic versant southwards to northern Panama; prefers moist friable soil, bur may be found in a variety of soil types.

2. An off-hand scale for size

Description: Body annuli ~ ave. 103 primary folds, ave. 62 secondary folds; is tailless; orbit is not roofed over by squamosal bone; body is thick; tentacle is located tentatively halfway between eye and nare on both sides of snout; absence of splenial teeth; overall coloration is a dorsum dark gray with a paler gray ventrum, jaw and tentacle markings; exhibits variable scalation in species, there are no subspecies.

Diet: Composed primarily of invertebrates that live in soils and under leaf litter such as worms, beetle pupae, termites, caterpillars and other insects; has been known to take very small vertebrates such as lizards, snakes and baby mice; is a sit and wait predator.

Habits: Unlike most amphibians, a hypaxial muscle does not aid in exhaling; to exhale, passive exhalation is achieved simply by opening the glottis allowing higher pressure lung air to exit into the lower pressured atmosphere; spends majority of time in burrows made in loose moist soils; prefers making their own burrows as opposed to using other animal digs; will come out and surface during warm gentle rains and crawl under ground litter; locomotion is achieved in wavy side-to-side accordion-like motions; is viviparous with female giving birth to 3-16 young always in May-June during the start-up of the rainy season; gestation is 11 months; in the first three months the fetus consumes egg yolk; in the final 8 months of fetal development the female releases a liquid nutrient from glands lining her egg ducts that the developing young consume with infant teeth; developing young also possess three branched gills to breathe while in the egg ducts.

Vulnerable 3.1 (IUCN)

1. Congo caecilian ~ H. squalostoma

Congo Caecilian
Family: Caeciliidae (Herpelidae)
Genus: Herpele
Species: Herpele squalostoma

Length: 40-60cm/15.-23.62in.
Weight: ~ 95-155g/ 3.35-5.47oz
Longevity: Maximum 18.5 yrs.

Name Origin: Herpele squalostoma ~ Herpetos originally Greek for 'skin'; with variant ele addition to refer to 'scales;' squalus is Latin for 'shark;' stoma is Latin for 'mouth;' together, all refers to 'scales in skin shark mouthed.'

2. H. squalostoma

Distribution: Is distributed in Africa from southeastern Nigeria through the Cameroon, the Congo, western Democratic Republic of Congo, southwestern Central African Republic, mainland Equatorial Guinea, Gabon, Bioko Island and presumably occurs in the Cabinda Enclave of Angola, although no species have been documented from there; prefers lowland rain forests with sandy soils adjacent to permanent bodies of water.

Description: Multiple ringed all down the body with annuli completely encircling dorsum and ventrum; possesses man dermal scales; mouth is large with unusually high number of splenial recurved teeth; has peculiar morphologies not found in other caecilians- possesses perforated stapes (stirrup-shaped small ear bone for high frequency detection), multiple small antotic (skullcase region) foramina (foramen- bone openings) and has no separate septomaxillae (septomaxilla- delicate bone associated with nare) and prefrontals [bone separating lacrimal (smallest facial bone) from frontal bones]; coloration is blue to light grayish purple; though study is ongoing to determine if is a single species, there are no current subspecies.

4. Profile of more blue specimen

3. Profile of more purple specimen

Diet: A generalist predator consuming fossorial invertebrates and occasional terrestrial invertebrates; diet may include invertebrate or small vertebrate carrion.

Habits: Life in the wilds is poorly known as majority of time is spent underground; may rarely frequent leaf litter; actual breeding is unknown, but are oviparous; appears to require moist soils; this caecilian is very primitive from other caecilians in representing the last line of an evolutionary lineage that first split off from the main branch over 95 mya; in this regard, the mammalians, man and mouse are phylogenetically more similar to one another than the Congo caecilian is to its closest relatives in the caecilian genus, Boulengerula.

1. Daniel's caecilian ~ G. danieli

Daniel's Caecilian
Family: Caecilidae (Indotyphlidae)
Genus: Gegenophis
Species: Gegeneophis danieli

Length: 150-195mm/5.9-7.7in
Weight: 4.2-8.3g/.15-.30oz
Longevity: Unknown

Name Origin: Gegeneophis danieli ~ Gegene is Greek for genea meaning 'race;' ophis is Greek meaning 'serpent;' danieli is a rendering for 'of Daniel;' all together refers to, ' Daniel's caecilian snake species.'

Distribution: Endemic to India in the Sindhudurg District of Maharashtra State within the Western Ghats; found in grassland patches surrounded by semi-evergreen forests.

Description: Except for G. nadkarnii, Daniel's caecilian differs from all other Gegeneophis species in possessing multiple dorsum secondary annuli (>50) in which most are complete midorsally encircling the ventrum; annuli are distinct displaying white edging; posterior to 35th primary annulus, there are few primay annulus on which the secondary annuli are incomplete middorsally; body shape is subcylindrical being dorsoventrally compressed; eyes are barely visible located under bone; rounded tongue is anteriorly unattached and separated by a groove from the gingivae (gums); nuchal grooves marked by whitish grooves; the cloacal opening is at the posterior terminus confirming no tail is present; overall coloration is dorsally a slate reddish gray to purple with a lighter colored ventrum; snout tip and lower jaw are pinkish; adult female body fat fluctuates in total mass, but not in length or body size; was just discovered in 2003 and there are no known subspecies.

2. 1. Body 2. Head 3. Rear

Diet: Bulk of food intake are subterranean worms (earthworms and grubworms); when not underground is under rock or ground litter during rains where it may consume invertebrate carrion such as dead insects.

Habits: As fairly newly discovered by hepretologist, Varad Giri, very little information is known in the knowledge of natural history and habits; Is strongly fossorial; when rains occur comes to surface; extensive burrowing aerates soils; perhaps mucoid secretions also play a role in chemically enriching soils; only one specimen was found and being female determined the species to be oviparous; eggs most likely are terrestrially laid where upon the hatchlings are fully developed.

Data Deficient (3.1) IUCN

1. Ringed cacilian ~ S. annulatus

Ringed Caecilian

Family: Caeciliidae (Siphonopidae)

Genus: Siphonops

Species: Siphonops annulatus

Length: 25-45cm/9.85-17.72in

Weight: ~ 89.5-143g/2.75-5.59oz

Longevity: environment dependent between 5-20 yrs.

Name Origin: Siphonops annulatus ~ Siphon (siphon pipe) and ops (face) are Greek and refers to tentacles on each side of face; annulatus is Latin meaning 'rings' in reference to body annuli; in describing these features tranliterates into tentacled face and body ringed caecilian.

Distribution: Dispersed throughout South America east of the Andes in populations found in Argentina, Bolivia, Brazil, Columbia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Suriname & Venezuela; is the most widespread of all caecilian species; appears to require both drier soils as well as humid in neotropical environments of forests and dry savannas.

Description: General aspect of the body is cylindrical and distinctly ringed from nape to vent appearing somewhat like a millipede without legs; diameter of body is 16th-17th the length of body; terminus after vent and head have no folds and are smooth skinned; tongue is surafce furrowed with vermiculiform (worm-like) sinkings with no protruberances; muzzle very short, thick & rounded; nare openings on very end and slightly more upward on snout; indentation below each eye appear as false nares; eyes are discenible embedded in skin; maxiillary and palatine teeth are strongly constructed, needle pointed and slightl recurved; 86-90 annular body folds equidistant from one another; cloaca is located at terminus end showing no tail; no apparaent dermal scalation (scutes); overall coloration is a navy blue or brilliant dark blue-gray with white edged annulus folds; there are no subspecies.

2. Head profile

Diet: Almost exclusively feeds on earthworms as it burrows through subsoil; on occasion takes other invertebrates such as fossorial arthropods and their larvae; is preyed on by snake predators such as Clelia clelia (the mussurana) and the semi fossorial Micrurus coralinus (tropical coral snake), with both being immune to venom of venomous snakes in which they both also consume.

Habits: In being nocturnal is most active at night; as in most caecilians, secretes skin toxins, but secretes a more potent toxin on the head and tail-end since these are the most apt areas to be grabbed by a potential predator; health is maintained by living in humid and dry soils; drier soils may help eliminate fungal growths; rarely takes to the surface; female is highly maternal in her parental investing; for reproduction consumes a lot of her energy and is labor intensive and as such, the female only reproduces ever two years; this caecilian is oviparous and lays a clutch containing 5-16 eggs; the female tends the eggs coiled around them for protection, added warmth and stabilization; once hatched the 75-135mm/2.95-5.31in neonates (infants/newborns) are altricial in that they require nourishment, but are incapable of movement on their own to acquire it; feed from the mother in two ways via the maternal skin and cloacal secretions; with 44 spoon-shaped and spiked teeth they practice maternal dermatophagy by rasping off the built-up outer skin of the mother and ingesting it; the other nutrient source is a supplementary cloacal exudation that the pups (young) imbide; the outer skin, rich in lipid fats & protein with the additional cloacal nutrients increases neonate growth rates on average per day .45% in length and 1.25% in weight mass; after attaining lengths beyond 150mm/5.91in, the pups leave the mother to forage and fend for themselves; throughout maternal care, female pales in coloration.

In the composite photograph to the left, a) is displaying the mother with neonates and still unhatched eggs; b) mother with altricial pups; c) pups collected around mother's cloaca sipping up the exuded liquid nutrient; d) enlarged view of a pups lower jaw and teeth; e) enlarged view of two spoon shaped infant dentary teeth. The final bottom three photographs are comparisons between the female's f) non brooding skin to the g) & h) thickened brooding skin after being stained.

1. Kirk's caecilian ~ S. kirkii

Kirk's Caecilian
Family: Scolecomorphidae
Genus: Scolecomorphis
Species: Scolecomorphis kirkii

Length: 215-463mm/ 8.47-18.23in (largest of Scolecomorhids)
Weight: ~ 130g/4.6oz
Longevity: ~ 14 yrs

Name origin: Scolecomorphus kirkii ~ Scole is derived from the Greek word, scolex meaning 'worm;' morphus is derived from the word, morphe meaning 'form' or 'shape;' kirkii is in honor of Sir John Kirk, a naturalist who was David Livingston's chief assistant during his second Zambezi expedition (1858-1863); all together refers to 'Kirk's wormed shape caecilian.'

Distribution: Habitat range is in eastern equatorial Africa in the countries of Malawi, Mozambique and the Ubena & Mahenge highlands and Udzungwa & Uluguru mountains of Tanzania; inhabits tropical rainforests in mountainous regions burrowed in humid soils or underneath jungle surface litter.

Description: Expresses dimorphism with the female being larger with primary annuli ranging from 140-152 in females and 130-142 in males; this female elongation with more vertebrae is an advantage in providing more space for developing fetuses; skin segmentation is reduced normally with only primary annulus; a single series of small teeth found in lower jaw; the mouth is subterminal; barely detectable eyes are roofed over by skin and bone; eyes are connected to tentacles; the flap shaped tentacles have an external subglobular base surrounded by a groove and a central opening through which the tentacles pass when being extended and retracted; tentacle apertures are positioned are positioned ventrolaterally from the snout below the nostrils and slightly angled to the anterior margin of the mouth; only caecilian genus where stapes are absent; epidermal scales are absent though rarely, vestigial scales are traceable in the posterior folds; there is no tail, where the bluntly rounded ventrally flattened terminus ends in a terminal shield without annuli; overall coloration is a dorsal lavender gray to purplish extending to the sides of the ventrum; ventral midsection is cream colored; tops and sides of head are a darker lavender shade than dorsum, no subspecies are recognized.

Diet: This caecilian is an active hunter consuming fossorial invertebrates, such as earthworms, grubworms and termites and terrestrial macro-invertebrates such as insects; it is common to find soil in their guts and whether this is intentionally taken in, or is remains from animals consumed such as earthworms, is not definite; is preyed upon by the fossorial burrowing snake, Atractaspis aterrima.

Habits: Kirk's caecilian is an active hunter and efficient burrowing predator, as has been observed tunneling while constantly extending its tentacles picking up chemical cues of the surrounding environment for prey; besides tunneling, this caecilian is probably the most active surface caecilian; when the tentacles are extended it pops the eyes out of socket and protrudes them beyond the head into the open; once the tentacles are fully retracted, the eyes go back in place; this ability insinuates light detection is more important than found in other caecilians; is viviparous with young retained in oviducts; fetuses possess fetal teeth as is found in all other

2. Head displaying left tentacle

viviparous caecilians, but they do not appear to practice intraoviductal feeding, as formerly pregnant S. kirkii females do not show any signs of oviduct scarring; a very large ovarian egg yolk and perhaps uterine milking that is stimulated by the fetal teeth is the prevailing thought for the source of this caecilian's fetal nourishment; if this strategy bears out to be true, it infers that fetal teeth were already in place and intraoviductal feeding was a later evolutionary edition as opposed to trending fetal teeth evolutionary development in accommodating more efficient oviduct scraping; the mother partakes in neonate parental care; whether it is for territorial rights or mating privileges, males show signs of aggressive bite marks that fit the pattern of male teeth.

1. Menon's caecilian ~ U. menoni

Menon's Caecilian
Family: Uraeotyphlidae
Genus: Uraeotyphlus
Species: Uraeotyphlus menoni

Length: 207-245mm/8.15-9.65in
Weight: ~ 90g/3.18oz
Longevity: Not established but estimated to be up to 14 yrs

Name Origin: Uraeotyphlus menoni ~ ur is a German prefix meaning, 'original' or 'primitive;' aeo is derived from the Latin word, aeon meaning a 'period of existence;' typhlus is derived from the Greek conjugation, typhlos referring to 'blind eye;' menoni is in honor of Dr. Sangeetha Menon from India; she is a zoologist, but is known more in the field of philosophy; altogether refers to, Menon's early blind caecilian.'

Distribution: As it is sometimes referred to as the Malabar caecilian, it is endemic to the southern portion of Malabar in the Western Ghats within the state of Kerala, India; its habitat seems to be limited to tropical and subtropical moist broadleaf forests.

Description: Both the head and terminus end taper; smooth tongue contours tapering jaws in being pointed; have a true tail with vertebrae; the tentacular openings are situated far in front of the eyes and below the nostrils (nares); eyes are distinct surrounded by a white ring; there is a total absence of tertiary annuli; trachea is expanded to accommodate larger surface area in respiratory gas exchange; there are between 165-172 annulary folds; dorsum coloration is violet to lavender that sometimes trends to being purplish with lighter groove segments for annuli; ventrum is paler in color; snout tip and tail tip are creme colored; no subspecies are recognized.

Diet: As it is strongly fossorial in habit, the bulk of consumption is probably subterranean invertebrates such as earthworms, termites and insect larvae.

2. Profile of tapered head & tail

Habits: Apparently this caecilian is restricted to really moist soils as it has only been found in areas receiving over 2000mm/78.74in of rainfall per year; has been found in humus-rich soils of tropical lowland forests and wetlands; it has also been reported from artificial terrestrial sites such as plantations and agriculture, but in that these sites were excavated from originally natural forests, soil chemistry will change and may be detrimental to this caecilian; morphology of tooth crowns is variable inter-specifically among caecilians and U. menoni possesses bicuspid teeth, which is indicative of prey choice as these teeth are utilized for prey grasping; as in all caecilians, insemination is internal; this species is oviparous laying a clutch of eggs in moist soil near permanent water; once hatched the larvae leave the nest site and enter the water appearing to be wholly aquatic during the larval stage.

Data Deficient (3.1) IUCN

1. Two-lined caecilian ~ R. bivittatum

Two-lined Caecilian
Family: Rhinatrematidae (Ichthyophiidae)
Genus: Rhinatrema
Species: Rhinatrema bivittatum

Length: Up to 210mm/8.27in
Weight: ~ 6.5-7.5g/.23-.25oz
Longevity: Unknown
Name Origin: Rhinatrema bivittatum ~ All nomenclature is Greek rooted; rhin refers to 'nose;' atremata means 'without holes' referring to lack of tentacular apertures on snout; bi is a reference for two; vittatus means 'striped;' altogether is: 'two-striped close-nosed' caecilian.

Distribution: From portions of northern South America in northern most part of the Amazon in Brazil, eastern most part of Guiana shield in northern French Guiana, Guyana and Suriname; found in lowland rainforests within these regions.

Description: Due to new DNA sequence data, the family Rhinatrematidae has just been deleted this year (2011) with its species being removed and listed under the family, Ichthyophiidae; Rhinatremata was a monotypic genus with R. bivittatum as the only species until in 2010, when a newly discovered species was found in R. shiv; rhinatrematids are the sister taxon of all caecilian species in being the most primitive branching off 215 mya; retain more ancestral features than other caecilian genera; R. bivittatum's skull is zygokrotaphic and is not fully fused lacking prefrontal & postfrontal bones and an opening between the squamosal and parietal bones; premaxillae and nasal bones are present as separate; eyes are distinct covered by clear tissue; tentacles are immediately adjacent to eyes and migrate anteriorly away from the eye; mouth is terminal on snout, hence the name beaked caecilian; has a distinct tail with vertebrae; have complete secondary annuli the full length of body and are orthoplicate (in same plane around body); overall coloration is dorsal brown with a splattering of yellow and a ventral majority yellow that extends to the sides, ergo the name two-striped; on both dorsum terminal ends is a large yellow spot; though at first Rhinatrema shiv was being considered a subspecies until it was determined to be distinct, there are no subspecies.

Diet: fossorial invertebrates such as earthworms and insect life stages and forest floor invertebrates suxh as insects and arachnids.

Habits: Is fossorial, but requires loose soil or humus for burrowing due to skull not being specialized for this activity; is probably the most active surface caecilian, therefore it is considered surface cryptic; usually haunts

1. Head profile

moist soils and leaf litter in foraging; are more nocturnal in habit and activity; occurs in remote habitat less likely affected by man; courtship, mating, though never observed, is considered to be terrestrial and underground; is oviparous with the female laying eggs in a terrestrial burrow near a water body source where she broods the clutch of eggs; once the eggs hatch the larvae exit the burrow and enter the water; larva is considered wholly aquatic, although there has been no observance, study or data on two-striped caecilian larval natural history; the population trend and status is not known, but is listed by IUCN as least concerned.

1. Koh Tao caecilian ~ I. kohtaoensis

Koh Tao Caecilian

Family: Ichthyophiidae

Genus: Ichthyophis

Species: Ichthyophis kohtaoensis

Length: 300-343mm/11.81-13.50in

Weight: 22-34g/.77-1.19oz

Longevity: Longest observed specimen lifespan ~ 7.3 yrs, but could be longer

Name Origin: Ichthyophis kohtaoensis ~ ichthys & ophis are Greek for 'fish' & 'serpent' respectively; kohtaoensis pertains to the island of Koh Tao in Thailand where was first classified as a species; all together is referenced as 'fish-like snake belonging to Koh Tao.'

Distribution: Known with certainty from Koh Tao, Thailand; the initial thought was to assign any yellow striped caecilian to this species nomenclature, but due to further taxa scrutiny, already the yellow striped caecilian from Yunan in China has been assigned a new species name, Ichthyophis bannanicus; ostensibly, the current attributed range is Indochina from the pennisular and mainland Thailand, Myanmar (Burma), Laos, Cambodia to Vietnam; populations are locally abundant in the Mekong Valley of Thailand; this caecilian prefers terrestrial microhabits of scrubs and forests near bodies of water; in these habits is within the soils during the dry season, while a greater percentage venture to the surface underneath ground litter during the rainy season.

2. Ventral side tapered tail

Description: Has a true but short tail with vertebrae extending past the cloaca (vent); females have a W-shaped wall at the posterior margin of the cloacal disc; mouth is not recessed underneath the head; possesses two sets of jaw muscles a unique feature found only in the family, Ichthyophiidae; primary annuli divided by secondary and tertiary grooves; eyes are visually distinguishable; tentacles immediately anterior to edge of eye; numerous scales evident in annular grooves and in some of the dorsal grooves of the nuchal area; skull is very zygokrotaphic descriptive; overall coloration is a dorsum dark gray and yellow ventrum extending to the lower to mid sides; lateral line systems composed of two elements, the neuromasts & ampullary organs, are developed during embryonic stage, while tentacles and yellow striping develop in the larva; for now, there are no assigned subspecies.

3. Embryo showing gills & lateral lines

4. Maternal care

5. Consuming a headless cricket carcass

Diet: Adults consume primarily subterranean invertebrates such as earthworms, insect larvae, termites, ants, oribatid & mesostigmatid mites and on occasion microhylid frogs; larvae consume strictly aquatic invertebrates, such as odonate larvae, water beetles, clam shrimp and will accept the carcass of dead invertebrates; moving from an aquatic larva to a metamorphosed terrestrial adult is a trophic position, but is superficial in both stages being generalist carnivores; predation of this caecilian by the red-tailed pipe snake, Cylindrophis ruffus is prevalent.

6. Consuming an insect pupa

Habits: Though primarily fossorial, during heavier rains can be common underneath surface litter; is primarily nocturnal being most active at night; female oviposits 32-58 eggs in a moist terrestrial burrow and remains with the egg clutch tending the eggs coiled around them until hatching; oviposition site is always near a body of water such as streams or pools; upon hatching, larvae exhibit positive phototaxis in utilizing light intensity to exit the burrow and find the nearby water body; well developed larval gills quickly degenerate and are lost within a day from time of hatching; larvae are not strictly aquatic, but are amphibious coming out of water by day to hide and retire under terrestrial retreats such as rocks, fallen logs and leaf litter at the edges of water; use negative phototaxis by day to locate dark recesses; by night they return to water to forage for aquatic insects, tadpoles, amphibian & fish eggs and terrestrial insects that have fallen into the water; lateral line system is highly developed in amphibious larvae and is utilized as mechano and electro-receptors to pick up physio ambient grade changes such as water pressure differentials and weak electrical fields produced by living organisms; terrestrial adults lose lateral line capabilities as it becomes functionless and disappears in adulthood; the muscle utilized for larva eye movement takes on a second role in adulthood as the retraction muscle for the tentacles; this caecilian appears to be adaptable to a degree in environmental changes.

1. Rubber eel ~ Typhlonectes natans

Rubber Eel

Family: Typhlonectidae

Genus: Typhlonectes

Species: Typhlonectes natans

Length: 45.7-55.9cm/18-22in; Female captives can reach up to 120cm/3.94ft.

Weight: 60-100g/2.12-3.53oz.

Longevity: 4-6 yrs.; Aquarists have reported captives living up to 20 yrs.

Name Origin: Typhlonectes natans ~ Ancient Greek - typhlos meaning 'blind' and nectes meaning 'swimmer;' natans is Latin for 'swimming.'

Distribution: Found in the South American countries Columbia and Venezuela, possibly in the Republic of Trinidad and Tobago; prefers subtropical to tropical bodies of water with mud bottoms in still waters of oxbow lakes and seasonally flooded river streams, shrub lands and low lying grasslands; will burrow into mud bottoms during dry season if caught in dried up seasonal water regions; waters warmer than 27.8 °C/82 °F will shorten life span.

Description: Are fully aquatic living exclusively in freshwater; cylindrical body is clad in dorsal mud gray with ventral coloration trending to lighter gray; some color morphs are more of a gray blue; eyes are covered over with skin; cloaca is located near the anterior end and has no tail; though has no true fins, possesses a fleshy ridge on dorsum anterior, there are no subspecies.

2. T. natans ~ blue morph

Diet: Aquatic invertebrates and their lavae and possibly scavenge dead animals lying on water beds; without true vision cannot actively pursue and capture fish.

3. T. natans ~ yawning

Habits: Will readily accept chunks of dead carcass in captivity; will bite into then roll and pull off bite size pieces; perhaps this habit is also practiced in the wild; captives will yawn often prior to shedding; breathing is primarily through cutaneous gas exchange though periodical trips to the surface to gulp air is necessary in adequate oxygen supply to the lungs; in gulping surface air, lungs are first emptied in a single extensive exhalation, then refilled with an inhalation series of 10-20 inspiratory buccal oscillations; are viviparous giving live birth to 3-7 fully formed young in water; until birth, fetal young feed off of the maternal oviduct; oviduct is hypertrophied with nutrients; young are born with long sheet-like gills that are released once the first gulps of surface breathing are performed immediately after birth; gills acting as a mammalian placenta for respiration probably are more functional during the long 40 minute plus period of labor by the mother than for after birth; if born in too deep a water, the young may drown before reaching the surface.

Salamanders:

Most authorities when explaining body features, express salamanders as superficially lizard-like in form and appearance and they are correct in that description. Even though the amphibian lineage is much older than the reptile line, lizards came long before the first salamander made the scene. Where the first known lizard fossils show up in the Triassic 220 mya, the first known salamander fossils are 55 million years younger. These oldest of salamander fossils were discovered in 1996 in China and Mongolia dug from Jurassic mudstones originating 165 mya.

Even though salamanders do not make-up the oldest specimens found in the fossil record, they are considered living fossils. The reason due is that essentially if you inspected an extant salamander pulled from underneath a rotting log or the fossil remains of one of the oldest salamanders known such as, Chunerpeton tianyiensis they would appear the same.

1. C. tianyiensis

Salamanders first evolved after the original Laurasia and Gondwana continents merged to form the super continent Pangea 200 mya. When the super continent began to split apart 180 mya, the salamander lineage hitched a ride on the Laurasian portion. By the time salamanders had evolved from their amphibian ancestory some 165 mya, Laurasia had indeed itself rifted into pieces that now form North America, Europe, northern Africa and northern Asia where current species reside. This is only where you will find today's salamander representatives and they all are related going back to the original salamanders that once shared a single habitat on the super continent Pangea.

With the above in mind, the Americas have by far more species of salamanders than the rest of the world combined with North America as the most prevalent of salamander species populations.

Because of distinct reproductive, physiological, morphological and biological shared traits between salamanders and frogs, it would appear that somewhere down their ancestral line laid a direct relationship. This theory has recently come to the forefront from a recent discovery of a 275 million year old fossil that was for all intents and purposes half salamander, half frog.

Living long before salamanders and frogs, Gerobatrachus hottoni was enjoying his days going after small insects in the Permian warm humid lands and taking to brackish ponds in eluding predation. First discovered in 1996 in scrubland near Waco, Texas by the late Smithsonian Institute paleontologist, Nicolas Hotton, the siltstone find languished in a Smithsonian basement collection until the mid 2000s when it was finally evaluated.

G. hottoni had teeth, but other than that resembled both the modern day salamander and frog groups. In frog characteristics, it had a developed tympanum, reduced vertebrae and a skull fashioned just like an anuran's in being broad while supported by long arching struts. In resembling salamanders, it had the same physiological body plan and shares in the fusion of specific ankle bones.

2. G. hottoni

This archaic creature may one day prove not to be the direct missing link to common day salamanders and frogs, but it is definitely a link-up to salamanders and frogs with solid inference that they do indeed share a common ancestry.

Of the approximate 350 salamander species worldwide, more than 180 are confined to the New World, or if preferred the Americas. That accounts for over half of the world's amphibian species. Northern South America, which was a part of Gondwana and not Laurasia does have a salamander population in the genus, Bolitoglossa. Belonging to the family Plethodontidae, which originated in North America, Bolitoglossa species most probably migrated to South America via the Central American land bridge, as Bolitoglossa species can be found from the U.S. down to Mexico, Central America and finally into northern and north central South America.

The trend now is to class all extinct salamanders under the order, Caudata, while putting all extant species under, Urodela.

Most salamander species are no more than 15.2cm/6in and range from the minute Thorius arboreus at 17mm/0.67in to the Chinese giant salamander, Andrias davidianus reaching lengths to 1.8m/5.9ft. Salamanders continually grow well beyond sexual maturity, so have to shed. Most, once shedding is complete, dispose of the dead skin by eating it.

With most requiring an aquatic stage, all salamanders are dependent on a moist or at least damp environment in being prone to desiccation. Even though the skin is enshrined in mucous secreting glands it does not retain water in extended arid conditions and is conducive to allowing moisture to diffuse not only inwards (internally), but outwards (externally) as well.

All larval stages possess gills for breathing and in permanently aquatic neotenic species gills are retained throughpout life as a larva. Given dosages of iodine or other thyroid hormones though, the larva will metamorphose into a terrestrial adult with developed lungs. It is thought that cooler temperatures, where most neoteny occurs, is the culprit in slowing or retarding maturation, as cooler temperatures inhibits the release of thyroxin and thus slows body growth. Neotenic salamanders can breed and produce offspring such as the axolotl. Gilled species will also utilize buccal pumping (gulping surface air) for respiraton. While some terrestrial species have lungs, most are lungless and breathe by cutaneous gas exchange through the skin and thin membranes in the mouth and throat.

3. Ensatinas nesting

The 'typical' life cycle of oviparous salamanders starts in a gelatinous mass of 1-450 eggs laid in an aquatic environment. From there, eggs hatch into larvae for 1-3 months with periods of metamorphism. Once the final adult stage is reached, the aquatic environment is abandoned for a multi-year terrestrial life. The adults return to water to lay eggs in water repeating the stages. The 'atypical' life cycle of oviparous salamanders is exhibited by plethodontids in terrestrially laying eggs in fashioned nests usually under rotting logs or damp floor litter with extended maternal care in protection from predation, the ridding of fungal growths and keeping the 10-30 egg clutch nestled. The 'exception' life cycle is carried out by a few species of closely related salamanders such as Salamandra atra that exhibit ovoviviparity in giving birth to larvae or completely metamorphosed juveniles appearing as miniature adults.

4. Vomerine teeth & nasal openings

Nare or nostril openings are located forward in the roof of the mouth leading into a small sac. The sacs contain a small amount of olfactory epithelium. The rest of the nasal system for smelling purposes is the vomeronasal organ that is a little sac lined with the bulk of olfactory epithelium tissue. Salamanders are the only amphibian that has the organ connected to the rest of the nasal system.

The teeth of salamanders are normally equal sized and evenly spaced and are used more as inducing a frictional surface when grasping rather than true grip, though some species exhibit small recurved teeth. The base of each tooth is fastened to the maxillary and vomerine bones as anchylosed (anchored) to the inner side of an external alveolar ridged plate and is said to be very similar in nature to fish as being ichthyic in characteristics. Plethodonts have an unusual cluster of small parasphenoid teeth running down the midline of the roof of the mouth. As an exception though, amphiumas have teeth that are intended to leave a nasty bite.

5. Maxillary & inner vomerine teeth

All terrestrial salamanders have four squat legs that drag the body along the ground, but since there is no auditory development, what's lacked in hearing, the salamander's body, jaws and limbs in close proximity to the ground, detects and picks up earth vibrations. In aquatic forms, vibratory frequencies are picked up by a lateral line system, a row of sensors located on the sides of the head.

6. Parasphenoid teeth

The fully aquatic forms have webbed feet with very short toes for clinging to slippery surfaces, while the terrestrial salamanders have virtually no webbing at all with longer toes for digging. The forelimbs grow faster than and longer than the hind limbs in all salamanders. Most salamanders have four toes on the forelimbs and five on the hind limbs. With the exception of just two related salamanders of the same genus, Onychodactylus fischeri and O. japonicus, all salamander toes have no claws.

The tails of permanently aquatic salamanders are compressed where terrestrial species are more or less rounded. Most salamanders have the ability to break off a part of the tail and regenerate the portion lost. The ability to regenerate tails or even other body parts is due to the adult's retention of embryonic blastema stem cells (for more detail see: 'Legs Anyone' under Quick Facts).

Most salamanders secrete skin neurotoxins that range chemically from mild alkaloid irritants to the highly toxic and lethal tetrodotoxin (TTX). TTX is the most potent nonprotein toxin known that blocks sodium channels keeping nerve signals from being transferred. Salamandra salamandra has the ability to squeeze the toxic glands squirting toxin for some distance, while the Spanish ribbed newt, Pleurodeles waltl has the ability to pierce its skin with its own ribs that become doused and tipped with skin toxin when passing through. For most, the 'unken position' is a defensive posture performed where the body is arched exposing the most concentrated body areas of secreted toxin. To dissuade predation, the toxins must be ingested or directly enter the bloodstream like through a wound. In topical contact, the toxin may be rinsed off with no ill effects.

Strictly carnivorous but depending on size, salamander diets consist of anything they can swallow whole, including other salamanders. Typical consumption for smaller species are aquatic invertebrates and their larvae, which includes aquatic crustaceans, leeches, earthworms, insects and snails. The larger ones in addition will take small vertebrates such as mice, snakes, frogs and fish.

In lungless salamanders, muscles surrounding the hyoid bone contract exerting pressure that shoots the bone out of the mouth along with the attached tongue. The tip of the tongue contains a sticky goo of mucus that adheres to the prey the catapulted tongue was directed towards. Once the targeted prey is snagged, pelvic muscles are contracted reeling in the hyoid bone, tongue and prey back into the mouth.

7. Dicamptodon eating a mouse

8. A Spring eating a Dusky

There are a wide array of courtship displays and fertilization is internal though without copulation. This is accomplished with male spermatophores that are gelatinous pyramidal structures that are capped with a packet of sperm. The male deposits this on the ground floor or water bed where then the female retrieves the sperm cap with her cloacal lips. As the sperm cap rests in the cloacal canal, the jellylike eggs are fertilized as they pass through the cloaca.

All newts are salamanders, but not all salamanders are newts. Newts are specialized salamanders that do not have slippery skin for it is rougher and drier. Essentially aquatic, there are some land forms such as Notophthalmus viridescens' red eft stage, where aquatic larvae turn into terrestrial efts remaining ashore for one-three years. After the eft stage the newts return to water transforming into aquatic adults.

9. Lungless salamander projecting tongue

Except in newts where they are indistinct, costal groove counts along the body sides in salamanders are good indicators in identifying and distinguishing different species from one another.

Depending on which classification scheme, there are eight, or if you want to include Dicamptodontidae and Rhyacotritonidae, then there are ten families listed in describing extant salamanders.

1. Two-toed amphiuma ~ A. means

Two-toed Amphiuma
Family: Amphiumidae
Genus: Amphiuma
Species: Amphiuma means

Length: 36.8-74cm/14.49-19.9in

Weight: 460-790g/1.01-1.74lbs

Longevity: Average ~ 17yrs, maximum ~ 27yrs

Name Origin: Amphiuma means ~ Amphi is Greek meaning 'on both sides' and uma is also Greek for 'pneuma breathe,' most probably in reference to misassumption that it breathed both water and air; describer never gave reason for means, such as in honor of an unspecified person, but most probably refers to the Greek word 'menos' meaning 'fierce' expounding on its lack of hesitancy to bite.

Distribution: All 3 amphiuma species only inhabit the southeast portion of the United States from Virginia to Texas with A. means' range covering the coastal plains from southeast Virginia through all of Florida westward to southeast Louisiana; habitat is in very still waters such as swamps, bayous, ponds, sloughs and permanent ditches preferably in acidic waters.

Description: This amphibian, along with the one-toed and three-toed amphiumas, with their eel-like and snakelike body features are commonly called congo eel or congo snake; Congo has nothing to do with the southeastern states; these common names arose during slavery in the U.S. when people assumed amphiumas were brought over from Africa during the slave trade period and were really eels or some exotic snake; A. means has two toes attached to very small vestigial limbs; larva and adult possess fully developed lateral lines

2. A. means, note 2 toes & lateral lines

3. A. tridactylum, 3 toes

along body and head; possesses mucous glands secreting a slime over skin that aids in desiccation and predation; larvae hatch with gills, but are soon lost replaced by a gill slit; transpires exchanges in gas through long, vascularized & fully septated lungs and though are submersible aquatic animals, relies on aerial-aquatic gas exchange, for will drown continually submersed in warm waters; eyes are lidless;

4. A. pholeter, 1 toe

have some of the most developed teeth for amphibians that are sharp and recurved and with strong jaws are used in capturing and holding onto prey and in defense; 57-60 costal grooves are present with 58 on average; tail is laterally compressed making up ~ 22.5% of total length; amphiumas are a paedomorphic salamanders in that the are sexually mature before physically mature; overall dorsum coloration is bluish black with ventrum side a shade lighter; protein analysis by horizontal starch-gel electrophoreisis verified that A. pholeter (one-toed) is much more distinct, distant and more primitive than the other two amphiumas; hence, A means & A. tridactylum are genetically very similar; there are no subspecies currently recognized for A. means unless the close relationship between the two-toed & three-toed amphiumas are reevaluated.

Diet: Is strictly carnivorous consuming invertebrates and vertebrates; as an opportunist carnivore, diet includes any animal it can capture such as aquatic and terrestrial larval invertebrates, snails, crawdads (crayfish), frogs, salamanders, small slow fish, smaller snakes including Regina rigida that has very few predators due to excretion of very foul musk, sirens where ranges overlap and small rodents frequenting water edges; with larger prey amphiumas have been observed constricting or at least wrapping their body around the trapped animal; predators are primarily snakes such as Farancia, Neroidea and the cottonmouth, Agkistrodon piscivorous along with larger wading birds.

Habits: Unlike most amphibians, with razor sharp teeth and strong snapping jaws accompanied with a nasty attitude, amphiumas can and inflict a wounding bite; Is nocturnal where it does most of its foraging for prey, while by day remains in self-dug water or shore burrows up to a meter deep, or in abandoned crawdad holes, mammal delves or may retreat to water bottoms in jumbled debris usually with head protruding; although considered aquatic, cannot totally rely on breathing underwater; as submersed, gill slits supply dissolved

5. Note egg connecting stems

oxygen (DO) to fully developed lungs, but is nonexponential; will drown within two days of submersion in water at or above 25 °C/77 °F; the cooler the water the longer it can stay underwater; therefore, amphiumas are considered obligate air breathers in that they are restricted to underwater breathing in having to come up for air; lungs are only fully ventilated when animal surfaces for atmospheric air; being the largest amphibian in their range, they may be the greater percentage of biomass in their home waters; although most amphibians are mute, amphiumas can emit a clear whistle in ambient air when disturbed; even though the vestigial limbs appear useless, do aid in slithering across terrain to gain access to water resources and does traverse terrestrially at times during rains; if necessary, during droughts, to manage desiccation, may remain in

6. female guarding egg clutch

aestavation mucoid chambers dug in receding muds and stay there until rains replenish water source; have been known to survive up to 3 years without eating while aestavating in a low metabolic state; with males testes enlarging and cloaca swelling between January & April, breeding begins in water during the Spring; male chooses female and instead of depositing the usual amphibian spertmatophore, conducts the closest thing to amphibian copulation and that is cloacal apposition; the male buts his cloaca up to the female's and while the cloacas are in a, let's say kissing embrace, he ejects sperm, thus injecting sperm into the female's cloaca where the eggs are then fertilized as she oviposits; female only makes a nesting site on land, but always near shore in damp terrain usually in mud; the nest is a hollowed out chamber usually under objects such as rocks or fallen logs, or may be just under surface of ground as a depression; the clutch consists of 40-200 eggs that the female will remain with, coiled around the clutch providing an incubation environment and protection until hatching, which is around 5 months from being laid; the 10 mm/0.39in eggs are laid in long strings connected by a stem that was formed by a rubbery outer coating that is also 10mm; as incubation progresses, the eggs darken turning from yellow, as yolk is consumed, to a cranberry coloration making the egg clutch appear as a clump of fruit; larvae, beginning life at 45-64mm/1.77-2.55in hatch with gills and immediately seek water; although threatened by pollution, habitat loss and dredging or muck removal, A. means is considered not threatened.

7. Note vestigial limbs

1. Mudpuppy ~ N. maculosus

Mudpuppy

Family: Necturidae

Genus: Necturus

Species: Necturus maculosus

Length: Ave~ 20-33cm/ 8-13in; can exceed 41cm/16in

Weight: 60-222g/2.12-7.83oz

Longevity: Average lifespan in wilds 11yrs; captives 20 yrs

Name Origin: Necturus maculosus ~ Necturus references 2 Greek words, nektos for 'swimming' and oura meaning 'tail'; maculosus is Latin meaning 'full of spots' in reference to somewhat spotty dorsum; altogether refers to 'spotted animal of aquatic existence.'

2. Note gills

Distribution: Range is from southeast Manitoba to southern Quebec down through the U.S. midwest to Missouri, then eastwards to North Carolina & Georgia and south to Mississippi; within range found in rivers, perennial streams, weed choked ponds and some large lakes.

Description: N. maculosus is known as the Common mudpuppy or waterdog as it is known in the southern states; retains brightly maroon colored gills throughout life in being neotenic; the ones found in colder waters with higher water oxygen levels have shorter gills than those found in the southern half of range; respiration is conducted with gills, cutaneous gas exchange and lungs; requiring all 3 methods of breathing and cannot rely on either 2 or less; head is flattened; tail is laterally compressed and short for overall size of animal; 4 toes are present on all 4 well developed flattened limbs in contrast to most salamanders that have 5 toes on hind limbs; male & female are very similar in appearance except that male's cloacae possesses 2 prominent papillae that directs backward and females do not; mucus glands secrete slime throughout the body; granular glands exude a toxin containing denatonium ions that are very bitter; gills appear maroon due to distal portion of gills being very filamentous containing many capillaries; lateral line systems aid in detection of water movement and pressure differentials; overall body coloration is a dorsal gray, rusty brown or blackish; black to bluish black spotting is random and may appear as many, even running into each other in forming stripes, or just a few simple blotches; ventrum side is generally a lighter gray while at times exhibiting a few bluish black spots; there are 3 recognized subspecies: Necturus maculosus maculosus, N. maculosus stictus & N. m. louisianensis.

Diet: As an opportunistic carnivore will take whatever is offered in the water, in which is a variety of aquatic organisms; the waterdog menu consists of aquatic larvae, aquatic insects, aquatic worms, snails, fish eggs, amphibians, small fish and carrion; most preferred food is crawdads (crayfish); on sides of mouth, lips interlock, allowing them to suck in their prey.

3. Eating favored prey

Habits: Is considered wholly aquatic and is rarely seen out of water; this salamander is very cold tolerant so does not hibernate, but instead retreats to deeper waters; in its colder range, has been seen swimming underneath ice; is one of the deepest underwater amphibians reaching depths of more than 30m/99ft; will inhabit murky or clear water; in the clearer, therefore more receiving of light, mudpuppies will appear darker and vice-versa, in murkier waters will tend to be lighter in color; with poor eyesight that is only best at light perception, depends more on its hearing, smelling and lateral line senses; prefers remaining on bottoms where as nocturnal uses its legs for walking the water beds in search of food during dark hours and remains hidden during the day on the bottoms under debris, rocks or sunken logs; prefers walking, but in alligator fashion, will sinusoidal tail swim with legs tucked to sides of body; in eating carrion, will readily take a hook baited with dead or decaying organisms, where usually the fisherman is ignorant of his totally harmless catch and feels he hooked a dangerous animal; females reach sexual maturity in 6 yrs; courtship takes place in the fall while in the southern most range is in winter; males join females in shallow water aggregations sheltered under rock; swimming in and around females, males then drop a 1cm/0.4in spermataphore from his swollen cloaca; female then picks it up with her cloaca and stores it in a special gland called spermatheca until spring; sperm fertilizes the eggs as she oviposits in spring months in an excavated nest cavity under rock or logs; the 18-180 (usually 60) unpigmented eggs are suspended by female to ceiling of nest cavity; she remains with and incubates the eggs until hatching around 40 days later from being laid; hatchlings are 22.5mm/0.89in in length live off the yolk until it is consumed and they measure around 36mm/1.42in.

1. Hellbender ~ C. alleganiensis

Hellbender
Family: Cryptobranchidae
Genus: Cryptobranchus
Species: Cryptobranchus alleganiensis

Length: 30.5-51cm/12-20in; record ~ 74cm/29.1in
Weight: 1.8-2.3kg/4-5lbs
Longevity: Estimation from length is 25 yrs in wilds; captive 29 yrs
Name Origin: Cryptobranchus alleganiensis ~ Both cryptos & branchus derived from Greek, kryptos meaning 'hidden' & branchion meaning 'gill' referring to 'lack of external gills'; alleganiensis pertains to the Allegheny Mountains; altogether refers to, 'gill-less salamander of the Allegheny Mountains.

Distribution: Range, including subspecies is Susquehanna River drainage from southeast New York down to Chesapeake Bay head in Maryland downwards to Georgia & northern Alabama, southwestward to extreme northeast Mississippi & into southern Illinois; isolated pocket of subspecies, C. a. bishopi in North Fork of White River in southeast Missouri & adjacent Arkansas into sections of Black River; essential habitat is clean rivers and larger streams with silt free rocky bottoms.

2. Another color form

3. Note undulating skin & side folds

Description: This large and unusually grotesque salamander was dubbed 'hellbender' by early day settlers in America due to the fact with its hellish wrinkled skin and flaps had to be a creature from Hell and was hellbent on returning to Hell; other vernacular common names are, 'snot otter', 'devil dog' & 'Allegheny alligator'; is the largest salamander in the Western Hemisphere and third largest in the world; dorsoventrally flattened head & body with thick skin folds on both sides of body; has loose undulating skin; possesses one gill slit on each side of head hidden by skin flaps; hefty tail is laterally compressed; beady eyes are located on top of head; limbs are stout ending in typical salamander arrangement of 4 toes on each forelimb and 5 on each hind limb; very little dimorphism is exhibited except in size with male being slightly smaller on average; skin is very slimy supported by mucoid gland secretions; color is variable ranging from dorsum brown to yellowish brown, olive to olive gray and light gray; black spotting or darker mottling may or may not appear; ventrum is always lighter and has fewer markings; 2 subspecies are recognized in C. a. alleganiensis & C. a. bishopi.

Diet: Although coexists with trout in cool rapid streams, is very rare for this salamander to prey on fish, but will in winter time take small fish when primary food is in hibernation; main food source consist of crawdads, aquatic larvae, worms & large insects and other stream invertebrates; in crowded populations has been observed to practice intraspecicific predation consuming its own eggs and larvae; it is thought that this is a form of population density control; smaller hellbenders are preyed upon by larger fish, snakes and turtles.

4. A daytime favored nook

Habits: Is fully aquatic and primarily nocturnal being most active in the first 2 hours of darkness, but normally never straying far from its picked territory; by day prefers to rest on or under the rocky bottoms with only the head protruding; is very territorial and will be aggressive toward other hellbender invaders trespassing into its favored domain; prefers waters no deeper than 3m/9.9ft; adult hellbenders are very dependent on fast flowing water and wrinkled skin for their unique respiration; though they possess a pair of large, long and transparent lungs, they are poorly vascularized and are not sufficient to carry out adequate respiration; therefore, the hellbender employs special cutaneous modifications, a large body form and a peculiar behavioral method that combined, confer to integument proficient respiratory capabilities; gill slits work nominally in aquatic gas exchange and hellbenders are not crafted for buccopharyngeal (gulping) respiration, so the bulk of gas exchange occurs in the side skin folds rich in capillaries exposed to water currents; the large size and flat shape exposes more surface area to flowing water and to ensure that, the animal will rock or sway in place which continually breaks the barrier layers between skin and water exposing the capillaries to oxygenated water and the water to spent gas in the blood held by the capillaries; an osmotic gas exchange occurs completing the respiratory cycle; essentially, the skin and in particular the skin flaps are one big gill; generally breeding takes place in

5. Note larva gills & yolk sac

August & September where their external fertilization is an exception for normal amphibian internal reproduction; male fashions out a nest site underneath rocks or river debris with the entrance facing downstream; then persuades or forces if necessary, gravid females to enter and lay their 150-200 eggs; several females may use one nest compiling their yellow eggs into one large jelly mass; one nest site may have upwards to 2,000 eggs; eggs from each female are rosary-like in strings; once laying is complete, the male ejects sperm from his swollen cloaca uniformly over the nest and hence, fertilizing the eggs; female will remain at nest site moving eggs to ensure adequate oxidation; male may possibly stand guard; larvae hatch after 62-75 days of incubation; for first couple months larvae retain yolk sac for nutrition; when incomplete metamorphosis is experienced after 18 months, larva lose lobed forelimbs and paddle shaped hind limbs for muscular legs & toes and the external gills are folded into body replaced with the permanent adult gill slit; feeding is done through asymmetrical suction where with flexible lips and jaws, one side of mouth is opened, while next moving bilateral elements of mandibles & hyoid separately creating an unidirectional flow of water into mouth; this completely harmless salamander has populations that are declining or disappearing altogether due to chytrid fungus and man's activities of damning, waterway siltation and pollution.

Near Threatened (3.1) IUCN

1. Texas Blind Salamander ~ E. rathobuni

Texas Blind Salamander

Family: Plethodontidae

Genus: Eurycea
Species: Eurycea rathbuni

Length: 13cm/5in total length (TL)

Weight: ~ 2.5g/.10oz

Longevity: Between 10-11yrs

Name Origin: Eurycea rathbuni ~ Eurycea is in reference to Greek mythology of 'Eurydice', the wife of Orpheus who he tried to rescue from Hades; rathbuni is in honor of Richard Rathbun, a naturalist who once headed the U.S. Commission of Fish & Fisheries.

Distribution: Strictly endemic to the Edwards Aquifer as restricted in distribution to the San Marcos, Texas area; entire life span spent in cavernous subterranean stream waters; has been collected 58m/191.4ft underground in limestone fault zones while drilling water wells.

2. Eurycea skull

Description: Eurycea rathbuni along with the other two localized subterranean salamanders, E. robustus & E. waterlooensis, were formerly listed under the genus Typhlomolge meaning, 'blind salamander'; due to phylogenetic mitochondrial relationships, the name placement was proposed in 1965 and instituted in 2000 keeping Typhlomolge as restricted to common ancestor of Eurycea lineage; E. rathbuni is an obligate troglobite (cave dwelling) salamander as confined to underground waters due to evolved specialized physiology and minima & maxima temperature dependencies; has completely lost pigmentation and eyes; eyes are covered over by skin and as usual in vestigial eyes, the eyelids were the skin membrane that covered over; the skull is posteriorly flattened and broad making the snout appear duckbillish; inherited from plethodontid ancestry that ventured into subterranean waters eons ago, the Texas blind salamander retains the family's feature of lunglessness, relying on external small gills for respiration; the legs are long and spindly; as typical in most salamanders the forelimbs end in 4 digits while

3. Note gills & unique snout

the hind limbs have 5; possesses 12 costal grooves; tail is laterally compressed, tapers at the tip and marginally finned forming the bulk of its swimming propulsion; this salamander remains neotenic and does not fully metamorphose into an adult, in which would be a death knell in its dark, watery environment; the smooth and unpigmented skin is translucent giving an overall coloration as white or pinkish white and reveals internal organs through the sides and belly; there are no recognized subspecies.

Diet: Is strictly carnivorous and sharing its deep, dark, watery, fragile ecosystem with 40 plus small aquatic invertebrates, is the top predator; invertebrates taken as prey are snails, worms, amphipods, daphnia, freshwater copepods, small decapods like cavernous blind crayfish & shrimp and any small outside dead or alive insect or crustacean that may enter into its environment; feeding habits are indiscriminate where cannibalism has even been observed; occasionally floods flush the salamander to the surface where it is vulnerable to fish and reptile predation.

Habits: Most probably evolving from brook salamanders, this salamander travels through underground aquifers by following submerged ledges; the finned tail, as long as the body & head, is very efficient in propelling the salamander through water; after swimming short distances will stop and spread its legs floating down to settle on the bottom; there, it forages most by combing the bottoms in search of anything edible; as it forages, it swings the head back & forth in attempts to pick up any water pressure differentials as sensed by the lateral line system; cavernous water system temperatures remain constant throughout the year at 21 °C/69.8 °F; through its underworld evolvement, this salamander has become

4. Note spindly legs

5. Another overall view

dependent upon consistent temperature and unresponsive to surface climate cycles; therefore with gravid females and juveniles found year around, breeding is not limited to any season occurring anytime throughout the year; eggs it appears require the maintained temperature range to properly develop; gravid females will oviposit anywhere from 13-39 eggs singly on the bottoms of stilled waters or where there is very little current; eggs may at times be laid as clustered, but only with 2 or 3 eggs; embryo development appears rapid as eggs hatch anywhere from 12-21 days from being laid; the Texas blind salamander, along with the other subterranean aquifer species are very susceptible to urban and agricultural run-off that can

6. 1 day old eggs

7. 11 day old embryos

8. Larvae hatching at 21 days

find its way into underground waters; these salamanders require clean cool filtered waters; development and human population growth has not only introduced groundwater pollution sources, but in the pumping and lowering of the aquifers to maintain the growth has greatly threatened blind salamanders; many areas, such as Wonder Cave where the Texas blind salamander was once common and collected, cannot currently be found; though the total population has been unknown due to the inaccessibility of subterranean environments, this and other concerns have listed this salamander as threatened.

Vulnerable (3.1) IUCN

1. Yonahlossee Salamander

Yonahlossee Salamander

Family: Plethodontidae

Genus: Plethodon
Species: Plethodon yonahlossee

Length: 11-22cm/4.33-8.6in total length (TL)

Weight: 4.5-4.9g/.16-.17oz

Longevity: Estimation ~ 4-6yrs

Name Origin: Plethodon yonahlossee ~ Pleth refers to the Greek word, plethore meaning 'fullness' and odon is also Greek meaning 'teeth'; yonahlossee is originally an Indian word for 'trail of the bear' but the name is derived from Yonahlossee Road where the salamander was first described in North Carolina on Grandfather Mountain; altogether the scientific name refers to the 'salamander with a mouthful of teeth from Yonahlossee Road'.

Distribution: Endemic to the U.S. found in deciduous montane woodlands of western North Carolina, the Blue Ridge Mountains of eastern Tennessee & southern Virginia; prefers by day to hide underneath forest floor debris, rotting logs and rocks under damp conditions in elevation between 430-790m/1410.8-2591.9ft.

2. Plethodon y. longricus

Description: Though a member of the 'glutinosus' salamander group and sympatric to P. amplus, P. cylindraceus, P. glutinosus, P. metcalfi & P. montanus to wit possible hybridization with P. glutinosus, the yonahlossee salamander is morphologically & biochemically distinct; in 1978 the crevice salamander, P. longricus was assigned to P. yonahlossee based on morphological & allozyme evidence, however some authorities still classify the two as distinct species, or at least P. logngricus as a subspecies due to crevice salamander's strict preference to rocky outcropping habitat around the proximity of Bat Cave, North Carolina; is the largest eastern N. American plethodontid with the female usually a centimeter longer than male; as a plethodontid is lungless and lays terrestrial eggs, as land plethodontids do, where hatchlings are fully formed exhibiting no metamorphic stage; typically there are 15-16 costal grooves; a visually obvious nasolabial groove is evident that aids in chemoreception; is a typical looking salamander characterized by a distinct brownish red to red dorsum, that is continuous on the backs of adults while more spotted and blotched on juveniles; red marking is in stark contrast to overall black background; rest of body including ventrum has white markings and white fleckings; except for continuing arguments on whether P. y. longricus is a true subspecies or is just a color variant form of P. yonahlossee, there are no other recognized subspecies.

3. P. yonahlossee juvenile

Diet: Insects and other small terrestrial invertebrates are taken including on a regular basis nematodes, mites, ticks, spiders, millipedes, centipedes and earthworms; secreting toxins from the base of the tail acts as a deterrent to bird and mammal predation, but snakes and larger frogs and crayfish will prey on this salamander.

Habits: In being nocturnal, comes out from underneath daytime hiding spots past dusk to forage; they do possess the ability to move quickly and will make a hasty retreat to hide under larger forest floor debris or into burrows; though comingles with other salamander species, aggressively defends small territories from other yonahlossee salamander trespassers; during colder months in their range, will migrate vertically downwards into burrows and remain there until the climate warms; may also aestivate during drought periods; both sexes reach maturity in their third year and it is believed courtship occurs in August as segregated couples are most abundantly found together during that time of year; reproduction takes place terrestrially; spermatogenesis occurs in the female once coming out from hibernation in the spring resulting from last year's courtship; spermatogenesis is the process in which the female receives sperm with half the

Plethodon spp. exhibiting maternal care

number of chromosomes (haploid) known as somatic cells; she carries the somatic cells throughout hibernation until the cells progress into diploid states in the spring becoming spermatoza and fertilizing the eggs; females are oviparous and depending on female's size will lay between 19-27 eggs in late August or early September; she oviposits eggs in terrestrial burrows or underground cavities and will remain with the clutch until hatching occurs; hatchlings are fully formed minatures of adults except in average overall coloration and sexual maturity; extensive logging has depleted populations as their home ranges are very small and they do not tend to migrate to more accommodating ranges.

6. A ventral view

5. A dorsal view

1. O'Donnell's salamander ~ B. odonnelli

O'Donnell's Salamander
Family: Plethodontidae
Genus: Bolitoglossa
Species: Bolitoglossa odonnelli

Length: 16.5cm/6.5in total length (TL)
Weight: ≤ 2g/.07oz
Longevity: 12-15yrs
Name Origin: Bolitoglossa odonnelli ~ Derived from Greek, bolitoglossa refers to 'webbed-feet'; odonnelli pertains to 'of O'Donnell' in reference to American naturalist, D. J. O'Donnell; all together infers, 'O'Donnell's web foot salamander'.

Distribution: Endemic to the 100-1200m/328.1-3937.1ft slopes of Alta Verapaz and the Montanas del Mico in Guatemala; range also extends into more lower lands of Honduras; ranges in heavily humid tropical montane cloud forest regions in plant leaf sheaths and along streams.

Description: Belonging to the tropical climbing salamander genus, Bolitoglossa two main features stand out; almost half of the species in the genus have webbed-feet while all have naso-labial grooves, wherein males have extended groove protuberances called cirri that aids in chemoreception; the webbing is an evolved feature of N. American climbing salamander plethodontids in living an arboreal existence; as plethodontids migrated southwards, natural selection evolved morphological changes in adapting from a temperate climate to a neotropical one; the webbing is an adaptation for climbing and not for swimming; webs are produced by

2. Note webbed feet & naso-labials

glandular growths in between foot digits; toes cannot be moved individually, but the foot as a whole moves; webbed feet create a suction by placing the ventral smooth pad of the webbed foot on a broad leaf or limb; in climbing vertically, the outside margins of the feet adhere to the substrate, while the center is slightly lifted creating suction aided by mucoid secretions and the wet environment; in addtion O'donnel's salamander possesses peg-like grasping structures much like those of tree frogs lining the ventral side of the webbed foot pad; tongue is free to project outwards in capturing prey; the protraction/retraction muscles are controlled by peripheral structures as opposed to the central nervous system; tissue membranes at the floor of the mouth are reduced while the sublingual fold is completely absent; therefore plays no role in the tongue's projectile capabilities; tongue appears mushroom-like in form hence the derived name mushroomtongued salamander; sexual dimorphism is evident in females being slightly larger, but in dental dimorphism, males have larger maxillary & premaxillary teeth; also overall dorsal coloration is a light charcoal black with cream to off-white striated double lateral patterning and a gray underside; there has been much parallel and convergent evolution in the genus Bolitoglossa; though may be conspecific (variant same species) with B. mexicana, it is not, nor does it have subspecies.

Diet: Consumes essentially small flying, climbing and terrestrial insects such as beetles, ants, flies and these insect larvae; other invertebrates include small spiders, centipedes, mites and millipedes; on rare occasions vegetation is consumed; primary predator is Thamnophis & Long-tailed false coral snakes in lower elevations.

Habits: Is a nocturnal generalist either actively seeking out prey or sits and waits to ambush unsuspecting prey passing by; prefers, whether resting or active spending most of its time in low overhanging branches; primarily arboreal, but will on occasion forage on the ground floor; mating takes place usually above the forest floor where the male deposits his spermatophore on a leaf, small branch or epiphyte where the female then picks up the sperm package with her cloacal lips and deposits it into her cloacal canal; Is oviparous, laying eggs terrestrially in

3. Note: prehensile tail

damp furrows or shallow burrows; female performs parental care with the traditional plethodontid curling around the egg clutch; hatchlings are fully formed when hatched; there are very few amphibians with prehensile tails and this salamander is one of them; the photo to the left is an exceptional shot taken by the Alaskan photographer, Todd Pierson of an O'donnell salamander utilizing its prehensile tail; due to habitat destruction of cloud forests where moist conditions are essential for this tropical climbing salamander, populations are declining and juveniles are becoming more numerous than adults.

Endangered (3.1) IUCN

1. Pacific worm salamander

Pacific Worm Salamander
Family: Plethodontidae
Genus: Oedipina
Species: Oedipina pacificensis

Length: total length (TL) ~ 11.95cm/4.70in; snout-vent length (SVL) 4.25cm/1.67in
Weight: ≤ 2g/.07oz
Longevity: Unknown; possibly up to 20yrs

Name Origin: Oedipina pacificensis ~ Oedipina references 'Oedipus' of Greek mythology who was a tragic king accidentally killing his father and unknowingly married his mother, an indirect reference to all the sister taxon species in the genus; pacificensis refers to 'of the Pacific'; altogether relates to the 'Oedipus salamander from the Pacific side.'

Distribution: Found on the Pacific slopes of southern Costa Rica and to a lesser extent in adjacent southwestern Panama between altitudes of 5-1200m/16.5-3,960ft; prefers consistent humid conditions in wooded ranges, but may be found in clearings such as abandoned banana plantations.

Description: Belonging to the worm salamander genus, Oedipina this salamander shares that genus' characteristics in being long and pencil thin with well developed eyes, short limbs & tiny syndactylus(webbed) feet; with over 50 caudal vertebrae, the tail makes up over 64% of total length; tail is rounded becoming compressed at the tip; females are slightly larger than males; head is moderately broad with snout bluntly rounded; though poorly defined, there is a depression at base of occiput (back of head); mouth has a distinct sublingual fold, 12-14 maxillary teeth, 4 premaxillary teeth, 8 vomer teeth and 112 same sized paravomerine teeth; body

2. Profile

has 19 to 20 costal grooves; overall coloration is a dark slate gray often with white markings where limbs meet body and displays white stripes located post-occular ending with white spots behind the eyelids; morphologically, this salamander cannot be distinguished from O. gracilis & O. uniformis, but DNA sequences bear out distinct and major differences between the 3 species; there are no subspecies recognized.

Diet: Small ground dwelling or burrowing arthropods such as insects and spiders; as a means of defense against predation, this salamander body flips.

Habits: This salamander is semi-fossorial, but occasions surface wet moss, tree buttresses and can be found underneath leaf litter & rotting logs; appears worm-like and slow, but is agile and quick; is nocturnal when most active in foraging, but can be found during daylight amid heavy rains; in being a plethodontid, is lungless relying on subcutaneous gas exchange through the skin and mucus membranes in the mouth and throat; they use their developed naso-labial grooves for chemoreception in detecting prey, predators and in finding a mate; the tongue is

3. A couple

~ 80% the length of the body and is used as a projectile supported by extendable bones to capture prey; extended and retracted in milliseconds; the tongue tip is designed for gripping; when not in use, it is rolled up and stored in the body cavity where the lungs once were; is an efficient burrower; though limbs are nimble, for digging the small appendages are not used but the small size keeps limbs from being a hindrance during burrowing by the blunt snout; being voiceless and with no means in making noise, males go through a complex courtship in attracting a mate; constant pursuit of a female suitor, ritualistic dancing and gentle caressing of the female by the male is performed; the male also partakes in making a small incision on the female's skin to introduce his mental gland pheromone secretions; this is conducted by carefully rubbing
the mental gland on the surgical wound; once the female has become enamored and receptive, the male lays his cone shaped spermatophore on the ground to be picked up by the female's cloaca where she will then store it for up to 2 months before utilizing it to fertilize her eggs; egg clutches or parental care have not been observed, but are assumed to be terrestrial in underground burrows with full development upon hatching as is the case with other plethodontids; though is not commonly found; stable populations appear to be maintained.

1. Eastern newt salamander

Eastern Newt Salamander
Family: Salamandridae
Genus: Notophthalmus
Species: Notophthalmus viridescens

Length: Up to 12.7cm/5in
Weight: Adult mean weight 2g/.07oz
Longevity: 12-15yrs
Name Origin: Notophthalmus viridescens ~ Both being Greek, Noto means 'mark' or 'spot', while ophtalmus means 'eye'; viridescens is Latin meaning 'slightly green'; altogether means 'common green color eye-spotted newt'.

Distribution: With losses in localized demographic ranges and annual variance in populations, this salamander's current distribution most likely resembles its historic distribution; among U.S. salamanders, this newt has a large extent in distribution second only to the tiger salamanders' (Ambystoma tigrinum) range; are found throughout the eastern portion of the U. S. from northern Minnesota, southeastern Wisconsin, eastern Iowa, Illinois, extreme central east and southeastern Kansas, eastern Oklahoma to Texas down to the Gulf of Mexico; historically prefers wooded and wet forests with permanent bodies of water; will colonize newly dammed beaver ponds.

2. Land dwelling red eft

Description: Newts have 3 life stages and this salamander is no exception; there is an aquatic larval stage, a juvenile terrestrial stage known as an eft, then finally as an adult back to an aquatic stage; larvae possess gills, are light to dark shades of brownish green and do not leave the pond from where they were hatched and have a compressed finned tail for swimming; larvae lose the gills going through various metamorphic stages until they transform into the terrestrial red eft with developed lungs; efts are brick red to orange red with dispersed black dots and a row of dorsolateral black edged red spots; eft size is 2.5-7.6cm/1-3in; red efts may roam land for more than 3 years before returning to a permanent body of water where it transforms into the adult relying more on subcutaneous breathing; sexual dimorphism is evident in adults where the aquatic male has enlarged hind limbs & feet, and develops black rough patches on the insides of hind limbs and on the bottom tips of the hind toes during the breeding season; adult coloration is a dorsal olive green to yellow brown with a pale yellow to yellow ventral side splattered with small black speckings; the black bordered red dots are maintained throughout all 3 stages; some populations will exhibit neoteny in skipping the eft stage going straight from larval form to adult remaining aquatic and retaining gills; There are 4 recognized subspecies that range from southern Canada and the Great Lakes eastwards all the way to Gulf and Atlantic coastal states; they are: N. v. viridescens, N. v. piaropicola, N. v. louisianensis & N. v. dorsalis.

Diet: All 3 stages are carnivorous; natural food for adults are aquatic animals they can swallow whole such as aquatic insects and their larvae, tiny molluscs, copepods, leeches, worms, fish & amphibian eggs, small fish and the larvae of amphibians; efts feed primarily on terrestrial arthropods and their larvae; larvae & adults can literally decimate mosquito populations by consuming large quantities of aquatic mosquito larvae; natural predators are in the aquatic stage certain are fish and turtles while on occasion water snakes, mudpuppies, sirens and bullfrogs who have developed some immunity through ingesting of the toxin secreted by this salamander; hog nose snakes occasionally consume efts; many animals feed on the eggs including other salamanders, even the adult eastern newts will cannabalize their eggs; digesting larvae have also been found in the pitchers of the carnivorous plant, Sarracenia purpurea; certain leeches will avoid latching onto eastern newts, but some species of leeches do appear to be a major source of aquatic adult eastern newt mortality.

Habits: All stages of the eastern newt possess the neurotoxin, tetrodotoxin, but in various units per concentrate; eggs have the least amount where the red eft stage has 10 times more the amount than the adult; for defense, the eft and adult stages will perform the unken reflex posture exposing the areas of the body with the most skin secretions to the foe; the neurotoxin blocks the transfer of nerve signals to muscles; all stages, at least in periods are active dawn, noon, dusk and night, so are diurnal, nocturnal and crepuscular; larvae prefer to remain near

3. Red newt larva

mud bottoms, or in muddy waters no deeper than 0.5m/1.65ft for camouflage & protection; they tend to amass and congregate together in shallows filled with vegetation where they forage; once the larva has transformed into an eft, usually under the darkness of night or during rains, it leaves the water in sequential waves seeking out moist sites in the upland woods in leaf litter, moss or underneath rotting wood; efts will avoid direct sunlight while foraging the forest floor; the red eft stage, dependent upon ambient conditions, lasts between 1-3 years plus and may travel far from their origins; efts seek out a temperature environment near the range of 26-28 °C/ 78.8-82.4 °F; efts hibernate during winter months; once conditions are right, efts will home in on their original homesite, enter the water and transform into the aquatic adult; aquatic adults prefer to pick a home range section in the body of water they're in; if during drought their homesite begins to dry up they will remove themselves to other sections of the water; once the rains come back and the body of water is refilled, they will return to their original section of

4. A terrestrial adult eastern newt

water; each individual recognizes its neighbors; aquatic adults do not hibernate and have been observed actively swimming underneath ice; they prefer ice free waters and will cluster and aggregate where the water is ice free; some aquatic adults will leave their watery environment to live terrestrially for a year or two mimicking the lifestyle of the eft; the land dwelling eastern newt must also hibernate during winter; the smooth skin will become more warty and dry; both the red eft and terrestrial adult have an uncanny ability to orientate and find their way back to their original home site through magnetoreception; biogenic magnetite within their body is a ferromagnetic material and works twofold in magnetic orientation and light wavelength distinctions; the material reveals to the salamander polarity based inclination differentials and also acting as a sun dependent compass, will allow the salamander to home in orientating to short and long wavelength levels in distinguishing the nanometer lengths of the light waves; this is a form of electromagnetism; all three stages use chemoreception for finding prey and

5. N. viridescens egg mass

alluding predators, while the aquatic adults further utilize it in finding mates; courtship, in late winter or early spring involves a dance ritual between the paired salamanders where it ends in amplexus with the male wrapping and gripping the females nuchal (neck) area with his enlarged hind legs all the while rubbing the chin of the female with his snout wafting pheromones to her nostrils and fanning his tail; this episode of the courtship can last for several hours, so occasionally the female will swim to the surface for air with the male still embraced; the behavior intensifies until the male suddenly releases his grip and positions himself in front of her undulating his body; if she is receptive still, she will press her snout against his body or tail where the male then releases his spermatophore pak; he then nudges her into position to encourage her to pick up the pak with her cloacal lips; if she is not receptive she will escape by swimming off; the female will lay anywhere from 200-400 eggs in a mass but singly wrapping each egg in some type of detritus material; most of the time the fertilized eggs are not laid all at once; once laid, she abandons them and does not present any parental care; the eggs hatch into the first larval stage 3-5 weeks from being laid; though most populations in the eastern newt's range are healthy, habitat destruction for the red eft and persistent droughts for the aquatic phases are concerns.

1. Emperor newt ~ T. shanjing

Emperor Newt
Family: Salamandridae
Genus: Tylototriton
Species: Tylototriton shanjing

Length: Up to 20cm/8in total length (TL)
Weight: ♂~8-13g/0.29-0.46oz ♀~10-31g/0.35-1.09oz
Longevity: May exceed 10yrs

Name Origin: Tylototriton shanjing ~ Both Greek, tylos means 'lump' in reference to enlarged warts and Triton was a minor sea god; shan & jing are both derived from Mandarin meaning 'mountain' & 'spirit' respectively.

Distribution: Exclusive to China and endemic in western Yunnan Province from the mountains along Nu, Lancang & Yuan Rivers; prefers elevations between 100-2500m/330-8200ft along slow or stagnant water bodies in subtropical, but cool forests and wet grass fields.

Description: This salamander is a rough skinned newt with distinct bony ridges on top of the head and down the spine; a row of 30 or so colored warts runs laterally down both dorsal sides; the anterior top of the vertebrate column and skull are especially thick; tail is laterally compressed; overall the body is black with a pigmented orange head, parotoid glands, vertebral ridge, the rows of body warts, limbs and tail; though patterns remain the same, the orange can highly vary in individuals to yellows or reds, while the body in individuals can

2. blue body

appear dark navy blue; the emperor newt is also called a crocodile newt; there are 4 species of crocodile newts, which are T. shanjing, T. verrucosus, T. kweichownensis & T. taliangensis; T. shanjing has just recently been removed from under T. verrucosus ( Himalayan newt) due to mitochondrial Cyt b gene analysis, but some scientists dispute this, feeling it was premature with very little analysis studies on the Himalayan newt; there are no recognized subspecies.

Diet: Is a carnivore in larval and adult stages, pursuing invertebrates such as aquatic & terrestrial insects, insect larvae, worms and snails; predation of eggs & larvae are other amphibians, fish and cannibalism has been observed; the most concerned predator to adults are humans where local culture uses the dried out skins for medicinal purposes.

3. Yellow-patterned individual

Habits: the genus, Tylototriton is the most primitive salamandrid and once had extended populations into Europe during Tertiary times, where now they are limited to Asia; is nocturnal and although adults have a well equipped tail for swimming, is terrestrial only going into the water to breed or flee predation; this salamander is very toxic containing enough alkaloids from one individual that could kill up to 7,500 mice with an intraperitoneal LD50 of 11.5mg/kg; alkaloids are water soluble skin secretions; alkaloids contain phospholipase that hydrolyzes phospholipids into fatty acids and are also both proteolytic active & trypsin inhibitive; skin toxin also serves newt as an anti-bacterial agent; certain water snake and birds of prey species seem to have overcome the noxious tasting and toxic secretions, as they will readily devour the adult newts; the rows of warts are where the toxic skin glands lie while the glands are pressed up against the spines of the ribs, the newt has the ability to accelerate secretions when threatened; while on land, the coloration is obvious, but is an aposematic warning that it is toxic; in water during breeding, which is the adults most vulnerable time to predation, the coloration does add to cryptic camouflage as overall coloration breaks up as refracted through a water medium, blending the newt into the colors of the pebble beds; the exceptionally thick bone of the skull

4. Red patterned individual

& vertebrae is also a deterrent to smaller predators; these newts rarely take prey from underwater; emperor newts are passive overall exhibiting nonaggression towards one another or predators; mating, breeding and laying of eggs occurs in water; in the wilds the breeding season occurs in the months from May to August; in slow moving waters the courtship begins with the male wrestling a female using his snout and thickened forearms in which he interlocks with the female's forearms and begins gently dragging her; this interactions stretches the course of hours due to attempts of the female becoming receptive; once female becomes responsive

5. An emperor newt larva

external mating occurs; the male drops up to 3 cone-shaped spermatophores and then nudges or drags the female so as to allow her cloaca to make contact with the spermatophores, whereupon she picks up the cone-shaped packets; the female then undergoes a 7-21 day ova development; once the eggs are ready for deposition, she will seek out a suitable spot for ovipositing; eggs are laid singly or in 10-15 egg strings attached to plants, rocks or any other submerged material; an average of 40-60 eggs, but up to a 100 have been observed; eggs are ~2mm/.08in; after 10-18 days the eggs hatch into 9mm/.35in long tadpole larvae and remain in shallow waters no more than 15.2cm/6in deep; larvae are very slow in developing taking up to 100-150 days to metamorphose into the adult; once the larva has reached 15mm/.59in it transforms into the adult form and leaves the water; supplanting wild populations for the pet trade, human medicinal consumption and habitat destruction have put this newt into the threatened listing.

Near Threatened (3.1) IUCN

1. Alpine newt ~ I. alpestris

Alpine Newt

Family: Salamandridae

Genus: Ichthyosaura

Species: Ichthyosaura alpestris

Length: ♂~9cm/3.5in; ♀~12cm/4.7in total length (TL)

Weight: 2-3g/.07-.11 oz

Longevity: To around 7yrs, but up to 10yrs & up to 20yrs is possible in some populations

Name Origin: Ichthyosaura alpestris ~ Ichthyos & sauros is Modern Latin from Greek meaning 'fish' & 'lizard' respectively; alpestris derived from the Latin word 'alpes' means belonging to the Alps; altogether refers to 'fish lizard of the alpine'.

Distribution: Main distribution center is in the middle of Europe radiating outwards to Denmark as the northern extreme and southeastward through Romania & Bulgaria via the Carpathian & Balkan Mountains downwards to northern Italy, Montenegro highlands and central Greece; an isolated population exists naturally on the Iberian Peninsula and in northern England from a successful 1930's introduction; prefers forested areas with plenty of aquatic habitats from lowlands up to the alpine belt line 3000m/9000ft in altitude.

2. A) Terrestrial female & B) male

Description: Formerly listed as Mesotriton alpestris and before then, Triturus alpestris; phylogeography of I. alpestris has recently been investigated using mtDNA, revealing five major clades; Clade A consists of populations from southeastern Serbia, originating in late Miocene; Serbian lineage is ancestral to a western and an eastern lineage, with a mid-Pliocene divergence; western lineage is divided into Clade B (Italy) and C (central Europe and Iberia); eastern lineage is divided into Clade D (southern Balkans) and E (central-northern Balkans); eastern clades seem to have been isolated in multiple refugia during glaciation cycles based on high sequence divergence; western clades are thought to have colonized central, western and northeastern Europe from a possible refugium in central Europe; analysis indicates paedomorphic lineages of I. alpestris appear to have evolved during early to mid-Pleistocene, likely in response to the ongoing climatic fluctuations; tails are laterally

3. Top: aquatic male & female below

compressed and are around the same length as body; vomerine teeth exist in 2 rows diverging widely posteriorly & converging anteriorly; stark dimorphism in breeding coloration with stockier female having larger body, head & tail; males possess a more swollen cloaca; in terrestrial phase skin is granular with male retaining a remnant mid dorsal crest; dorsum is from dark grayish brown or dark olive to black; ventrum is always a shade of orange; in the breeding and aquatic phase skin becomes smooth; males exhibit dorsal blue coloring with stippled black & white flanks and a blue stripe at the edge of the orange ventrum; dorsal crest is more pronounced; female in the breeding & water phase exhibit a cryptic water camouflage coloration of overall mottled gray-green-brown exhibiting weak dorsum spotting and an orange ventrum; there are 10 recognized subspecies; they are: I. a. alpestris, I. a. apuanus, I. a. cyreni, I. a. inexpectatus, I. a. lacusnigri, I. a. montenegrinus, I. a. piperianus, I. a. reiseri, I. a. serdarus & I. a. veluchiensis.

Diet: Most invertebrates such as flying, terrestrial & aquatic insects, including their larvae are taken as well as small spiders, batrachian eggs & tadpoles, woodlice, slugs and snails; there are a host of predators including water & shore birds, owls, frogs, large predaceous aquatic insects such as water boatmen and insects with large aquatic larvae such as dragonflies.

4. Aquatic male showing off regalia

Habits: Is nocturnal coming out of its hiding place at night to slowly and methodically hunt for prey while in the terrestrial stage, though can show short acceleration in capturing prey or fleeing predators; in the aquatic phase, is active nocturnally and diurnally and without hesitation, quickly pursues prey; in aquatic phase is a very efficient swimmer and in cold waters may spend several minutes in voluntary apnea; collects dioxygene dissolved in water through subcutaneous skin gas exchange breathing, but will still maintain surface gulping; this salamander relies on sight and smell more so than most other salamanders; slowing down their rate of metabolism, they hibernate during winter months sheltered in rock cavities, underground tunnels or under wood heaps; when warmer weather comes in the spring, the terrestrial forms come out of hibernation and begin trekking to familiar

5. Larva

stagnant or slow shallow spawning pools; once there, they molt loosing their granular skin and transform into their aquatic breeding form; in attracting females in water, the male constantly posts himself in front of the female fanning his vibrant tail; once the effect is functional in submitting the female and she then begins following him, the male drops his spermatophore; once the female spots it, she proceeds to picking it up with her cloaca; these ballets continue for several weeks; at the end of courtship, the female now begins to pick out a favored spot of watered vegetation to begin laying her fertilized eggs one by one; the eggs are sticky adhering to the plants where with her legs she folds the plants around the eggs in hiding them from predation; once she has laid all her 70-190 transparent 2mm/.08in in diameter eggs, no further parental care is taken; depending on water temperature, in 1-2 weeks the eggs hatch into tiny 1cm/.40in larva that have no jaws or limbs, but a fully developed tail, gills and most of the time eyes; at this larval stage, hiding, resting and fleeing is the main activity; once larvae metamorphose into the second stage, a mobile neck and jaws have formed and begin seeking out nourishment in pursuing tiny insects, larvae and daphnia; anything that moves near them they will pounce; as larvae metamorphose, a dorsal fin down the back & tail appears, the forelimbs first appear, then finally the hind limbs; once the gills are lost, they have fully transformed into miniature adults, then leave the water for a terrestrial life until maturing into reproductive adults 2-3yrs later; some late hatched larvae have the ability to hibernate and sometimes neoteny occurs skipping the terrestrial phase; chemical pollution of wetlands and habitat destruction due to development have extirpated these newts from regional localities.

1. Ringed salamander ~ A. annulatum

Ringed Salamander

Family: Ambystomatidae

Genus: Ambystoma

Species: Ambystoma annulatum

Length: 14-24cm/5.5-9.4in snout-vent length (SVL); ave: 28cm/11in total length (TL)

Weight: 13g-0.46oz

Longevity: 4-6 yrs; possibly under good conditions up to 10yrs

Name Origin: Ambystoma annualtum ~ Anabystoma is New Latin meaning 'to cram into the mouth' with no understanding to reference; annulatum is Latin meaning 'furnished with rings' in reference to salamander's color ringed dorsum; altogether means, 'ringed salamander with a mouthful'.

Distribution: Endemic in the highlands of the Ozark & Ouachita Mountains along the borders of Arkansas, Missouri and Oklahoma; a small band has also been discovered in southwestern Illinois; populations in the northeastern portion of range (central Missouri) have least variability in mitochondrial DNA components from other populations, suggesting the northern populations are the most recently established from southern groups migrating northwards; prefers mixed hardwood-pine forested damp areas where temporary bodies of water occur in shallow holding ponds, trenches, ditches or seasonal creeks.

Description: Overall the body is well-rounded and slender in girth; the head is small with a depressed snout; tail is long and rounded, but slightly compressed; possesses vomerine teeth arrayed in 2 short series located behind the choanae (paired openings between nasal cavity & nasopharnyx); teeth are blunted & small; this species normally has 15 costal grooves; there are no sexual dimorphic characteristics and the species is considered monomorphic; typically, a

2. Variable dorsal ring coloration

brief light colored bar exists between the eyes and may extend below the eyes as pointed diagonally posterior; dorsal color of adults is a very dark gray to darkish brown with contrasting rings that vary in color from lime green to yellowish to whitish; dorsal ring coloration can even vary on the same individual; rings may also be broken up and blotch marked; ventral & limb coloration is usually gray to dull yellowish mottled with lighter spotting; there are no recognized subspecies.

Diet: Adults are vermivores (eats worms) consuming terrestrial & aquatic worms and grub worms; also eats other terrestrial very small amphibians, insects, non-insect terrestrial arthropods, land snails and mollusks; larvae consume cladocerans (water fleas), ostracods, copepods, small aquatic insects, aquatic dipteran larvae, dragonfly & damselfly nymphs and are obligatory oophagous with larger metamorphosed advanced larvae eating smaller first stage larvae or any remaining eggs; there is a long list of predators that prey on this salamander including vertebrates such as snakes, owls, shrews, skunks, raccoons and opossums.

Habits: The ringed salamander belongs to the mole salamander group due to leading a fossorial lifestyle; it spends most of its time in its own tunneling systems or in other animal burrows; at

3. A. annulatum ~ note small head

times can be found underneath objects on the forest floor such as leaf litter, or hidden under rocks and rotting logs; may be a voracious eater, but apparently does not consume food too often as prey studies conducted in examining stomach contents 90% or more had empty stomachs; is nocturnal when it is most active; when usually encountered is in early fall when colder rains initiate a stimulus to breed; once September rains occur this salamander begins to locally migrate in numbers on land to temporary water sites filled with rain water; migration to water sites are usually no more than several hundred meters from than terrestrial non-breeding sites to the temporary waters; from 10 to several thousand individuals make the migration to a single water site; males arrive first, entering water to wait on females' arrival; once females arrive and enter water, a frenzy erupts with individuals engaged in mass courtship; males may deposit their wide base spermatophore on the water bottom, on a submerged plant or even on each other, male or female; once a gravid female has picked up a spermatophore with her cloaca, thus fertilizing her eggs internally; she lays several clusters of up to 50 eggs in each on the water bottom or submerged vegetation; laying eggs in temporary water has advantage of no fish predation, but also has disadvantages of water source drying up and hypoxia due to lack of oxygen in stagnant shallow water; eggs appear to have solved these 2 dilemmas; during drought, eggs that fall into mud cracks or under the shade of overhanging plants can temporarily survive in dormancy up to 20days until the next rains come; under hypoxic conditions, the surface area of egg capsules increases oxygenation conductance and enhanced oxygen transport, thus increasing survival rates; female oviposits eggs on courtship night up to 2 nights afterwards; after 2 nights most individuals leave water and remain on surface up to a week befroe going subterranean; eggs hatch in 2-4 weeks from time of oviposition; larvae reach maturity at 48mm/1.9in 6-8.5 months from hatching; depending on ambient conditions,

4. Juvenile just lost gills

larvae complete final metmorphism from mid April to July in losing the dorsal fin and gills, while gaining strong limbs; primarily on rainy nights, the newly transformed juvenile will leave the water in search of underground shelter; juveniles take 2-3 yrs to become reproductively mature; both adults and larvae hibernate and aestivate; one of the major mortality rates is roads that traverse across ancestral migration paths where potentially hundreds if not thousands can be ran over.

1. Olm ~ Proteus anguinus

Olm

Family: Proteidae

Genus: Proteus

Species: Proteus anguinus

Length: 30cm/0.9-12in juvenile-adult total length (TL)

Weight: 2-150g/0.07-5.3oz juvenile-adult

Longevity: In wilds, average: 58-68.5yrs; maximum can reach over 100yrs

Name Origin: Proteus anguinus ~ In early Greek mythology, Proteus was a sea-god, whose name suggests 'the first'; anguinus comes originally from the Proto-Indo-European word, anguis meaing 'snake' and later modified in Latin to, anguinus meaning 'snake', 'serpent' or 'dragon'; altogether refers to the, 'primeval dragon'.

Distribution: Endemic to subterranean waters in the Dinaric Karst aquifer system of southern Europe; through limestone faults and caverns underground waters are fed by regional lakes and the Soca & Isonzo river waters; range includes northeastern Italy amid Trieste, southern Slovenia, southwestern Croatia, Bosnia & Herzegovina; the olm inhabits subterranean waters that flow underground through this extensive limestone region.

Description: A true troglobite, the olm is totally adapted to a dark aquatic existence and appears to share heterochrony (developmental changes) characteristics with another true troglobite salamander (for reference see: Texas blind salamander) such as a dispropotionately dorsolateral elongated head, frail limbs & lack of skin coloration; the olm is also neotenic with no clear metamorphosis maintaining the juvenile's external gills and compressed & finned tail throughout adulthood; eyes are visible in larvae, but are undeveloped and sealed over by skin by the time adulthood is reached; 3 pink feathery gills exist on each side of head;

2. Note similar head to E. rathbuni

forelimbs bear 3 toes while hind legs have 2 attached to a very elongate tube-shaped body; rudimentary lungs are semifunctional for respiratory breathing in hypoxic conditions; sexual dimorphism is evident in size with females being larger and size & shape of male's larger cloaca during breeding; are not true albinos in still possessing dormant pigmentation from original ancestry and will darken if exposed to sunlight; some juveniles will exhibit melanin spotting; skin is translucent white and reveals contours of internal organs on ventrum; due to translucent circulatory system can give a pink or peach tone to skin; olm is Germanic Latin originally referring to white people of central & northern Europe, thus this salamander is also called the 'human fish'; there is one recognized subspecies in the black olm, P. a. parkelj possessing active melanism; the black olm, found only in Bela Krajina of southeast Slovenia, visits surface waters retaining its pigmentation & eyesight.

3. Subspecies: P. a. parkelj

Diet: Is strictly carnivorous in all stages of life consuming invertebrates and their larvae, but mainly aquatic insect larvae that has been washed into cavernous waters of Diptera, Ephemeroptera, Plecoptera & Trichoptera insect species; also when available will consume amphipods, molluscs & worms; predation in subterranean waters are few, but is preyed upon by fish, toads and on rare occasion from cavern infestation of rodents.

Habits: This salamander spends its entire life in an aquatic totally dark environment; when active is solitary, while at rest is gregarious aggregating in water's stone bottom cracks or underneath rocks; for protection from invasive diseases and ectoparasites, the stratum mucosum (outermost skin cell layer) secretes a thin layer of mucous enveloping the body; of interest in comparison to surface dwelling aquatic neotenic salamanders, the olm in utilizing a lower metabolic rate consumes less oxygen in exerting same activity; when hypoxic conditions exist in stagnant underground pools, during low water levels, the olm can utilize the vestigial lungs to accommodate oxygen intake by breathing in surface air; though eyesight is only functional in detection of lightness/darkness, the olm has an array of receptors & sensors for a

4. Dorsal view

subterranean lifestyle; through chemoreception, is able to detect very low concentrations of organic material in water; though reduced, pineal organ affects skin pigmentation making skin cells photosensitive; located on inner surface of nasal cavity & in Jacobson's organ, the nasal epithelium is much thicker than found in other amphibians producing a very enhanced sense of smell; taste buds are located in mucous epithelium of mouth with most embedded in upper side of tongue and on entrance to gill cavities; buds in oral cavity are used for taste while those in gill cavities sense water's chemical compositions; mechanoreceptors, such as sensory epithelia of inner ear are specifically differentiated enabling the animal to receive & interpret sound waves traversing through water; as well as ground vibration; hearing capabilities underwater is evident in localizing potential prey; through ampullary organs located on the head, the olm is able to detect and register weak electric fields produced by organisms; there is also a geomagnetic sense capability in making use of the earth's geomagnetic field to orientate itself; the olm is a superb swimmer; with its short finned tail, undulating seprtentine body and limbs tucked against the body, it can swim up to 8kph/5mph; in subterranean waters, temperature is

5. Ventral view

constant so climatic variability as on the surface, give no clues to breeding cycles; therefore reproduction for the olm is year round; males become territorial and aggresive towards one another when breeding; during courtship, when a female enters a male's territory, he performs tail fanning with her sensing resultant waves picking up vibratory stimuli & odor cues; in responding, she approaches male stimulating him by rubbing her snout on his cloaca to turn around to her posteriorly and deposit a spermatophore on the water's bedrock; the male then stimulates the female by rubbing his snout over her cloaca; passing over the spermatophore, the female will then pick it up by drawing it into her cloacal cavity, thus fertilizing her eggs internally; courtship may be repeated several times in a period of hours; once courtship is final, the female leaves the male's territory in search of a suitable nesting ground and arranges a simple nest out of debris litter from the water's floor; after 2-3 days from courtship, she oviposits up to 70 eggs over the course of 25 days; eggs are guarded by the female until hatching; though temperature is constant temperature range does effect amount of time for incubation; egg development takes anywhere from 182 days to 86 days with colder ambient temperatures taking the longest and warmer the shortest; eggs double in size through water uptake; olm larvae take on the appearance of adults after a few months old and take up to 10-15 yrs in becoming sexually mature; subterranean waters olms live in is not a closed ecology and is dependent and susceptible to surface waters; if polluted waters inundate underground water, the highly specialized olm is endangered; over harvesting for the pet trade has also threatened this salamander.

Vulnerable (3.1) IUCN

The following last three salamander groups under the families: Dicamptodontidae, Rhyacotritonidae and Hynobiidae were not listed in the 'DISCUSSION' section under the outlined classification scheme. Not long ago, Dicamptodontidae and Rhyacotritonidae were removed from Ambystomatidae by classifiers listing both under Dicamptodontidae. Even more recently, rhyacotritonids were taken from dicamptodontids, receiving their own family title as Rhyacotritonide, while Hynobiidae was a short while ago delisted from the family, Cryptobranchidae. Dicamptodontids and rhyacotritonids are monotypic in genera, while hynobiids are listed under nine separate genera. All three families are still considered as sister taxons to their former family groups, but due to new genetic analysis and unique morphologies, the trend is now to list them under their own family.

1. California giant salamander

California Giant Salamander
Family: Dicamptodontidae
Genus: Dicamptodon
Species: Dicamptodon ensatus

Length: 6.25-17cm/2.5-6.8in snout-vent length (SVL);17-30.5cm/6.7-12in total length (TL)
Weight: Mean average: 20.8-24g/0.73-0.85oz
Longevity: Statistical life span is 6-10yrs; Maximum longevity: 17yrs
Name Origin: Dicamptodon ensatus ~ Di meaning 'two', kamptos meaning 'curved' & odon meaning 'teeth' are all of Greek origin; ensatus is Latin for 'sword-shaped'; altogether, refers to 'recurved pointed teeth salamander'.

Distribution: Endemic to U.S. western coastline from northern California to the extreme southern end of British Columbia covering only .03% of British Columbia's total land mass; in its range, inhabits wet & cold coastal temperate forests from sea level to 909.1m/3,000ft near clear fast flowing waters and seepages, but also at times near lakes and ponds; does not occur anywhere east of San Fransico Bay.

Description: Formerly known as the Pacific giant salamander where this colloquial name tag is now reserved for the Dicamptodontidae family and Dicamptodon genus; is one of the largest terrestrial salamanders in N. America; limbs are stout & supportive; possesses the typical salamander digits of 4 toes on the forelimbs & 5 on each hind limb; the California giant has a broad head with a shovel-like snout and a distinct gular fold (fold of skin across throat); tail is

2. Head & gills of neotenic form

laterally compressed and 40% of TL; large eyes have black pupils with brass-flecked irises; overall dorsal body coloration is a rust reddish brown with irregular marbled darker brown patterns; ventral side is egg-white or a very dull yellow and is normally unmarked; remaining aquatic, neotenic adults are common retaining gills and are a general brown with slightly darker brown markings; young larvae are streamlined for fast flowing water.

Diet: all phases are carnivorous predators; terrestrial form consumes insects, worms, snails ( in particular banana slugs which are common in their range), small frogs, small snakes & small rodents such as mice & shrews; neoteny form consumes aquatic insects & their larvae, other small amphibians & their larvae, small fish and small water snakes; larvae aggressively feed on any aquatic invertebrate that swims by and can be fitted into the mouth; predators of this salamander are carnivorous or omnivorous small mammals such as river otters,weasels, raccoons & opossums, large snakes & birds.

3. Consuming a banana slug

Habits: The California giant salamander as terrestrial and in its neotenic form, is elusive hiding underneath objects such as stones when at rest mostly during daylight; although it is primarily nocturnal, it will expend actively during wet daylight; though it is least active during the cold winter months, it does not hibernate; with hardened toe digits is a capable digger and climber in pursuing prey & shelter; directly sunlit areas are avoided preferring to range in damp areas to combat dessication of the skin; this is an aggressive animal as far as salamanders go and can inflict a painful bite with its strong jaws & tiny sharp recurved teeth; aggressive towards potential predators & their own, they will thrash the tail, bite and from glands located at the top base of the tail secrete foul tasting chemicals; neotenic adults are found throughout all populations, while at times may outnumber the terrestrial forms in a population density; is one of the few salamanders that can vocalize emitting a low barking-like sound when alarmed or threatened; breeding occurs from spring to

4. Note size to Ensatina escholtzii

autumn with terrestrial forms returning to suitable creeks and streams to breed congregating under submerged stones & logs; courtship ritual is not known, but fertilization is internal; after her eggs are fertilized, the female seeks out a suitable nest site under submerged objects or crevices; once 70-200 eggs are oviposited singly or in clumps, female, without eating, will relentlessly for 7 months protect the developing eggs from any intruders including males who will eat her eggs if given a chance; when the transparent eggs hatch, the young larvae remain at the nest site for 2-4 months; during this period, larvae do not feed surviving off yolk; due to long gestation period, females are biennial in reproducing every other year; females do not reach sexual maturity until 5-6 years old; young larvae remain in the stillest waters near shoreline, while the older venture into the main stream; to fully transform from larval stages to terrestrial adult takes 18 months; due to slow reproduction, habitat alteration and the building of roads near the streams these salamanders migrate to & from, the species is becoming threatened.

Near Threatened (3.1) IUCN

1. Olympic salamander ~ R. olympicus

Olympic Salamander
Family: Rhyacotritonidae
Genus: Rhyacotriton
Species: Rhyacotriton olympicus

Length: This small salamander never exceeds 10cm/3.9in total length (TL)
Weight: ~ 1.75-2.5g/.06-.09oz
Longevity: Not recorded, but due to cold environs slowing metabolism, >10yrs is likely
Name Origin: Rhyacotriton olympicus ~ Rhyakos is Greek for 'stream' while Triton was a Greek sea god; olymipcus refers to belonging to Mount Olympus; altogether references the, 'stream god salamander of Mount Olympus'.

Distribution: Limited in range to the Olympic Peninsula of northwestern Washington state; a prerequisite habitat of cold & wet environs of silt-sediment free streams in old mature forests is necessary for sustenance; this torrent salamander is associated with cold, clear flowing waters typically found on steep slopes, seeps, springs & waterfalls; is most abundant in or around splash zones of falling or cascading water.

Description: This small salamander's defined head has a short snout with large eyes; the horizontal diameter of the eyes are equal to or greater than distance from anterior of eye to tip of snout; dimorphism is prevalent with females slightly larger while males possess a prominent

2. Ventrum

square-edged lobe just behind cloaca; breathes primarily through subcutaneous gas exchange; lungs are reduced, but still functional; overall coloration is medium-dark brown interspersed with white flecking and darker mottling on the laterally compressed tail; ventrum is bright orange brown to yellow usually with a few to numerous black spotting between throat and underside of chin; tiny larvae have very short external gills, flattened bodies and fins limited to the tail; larval coloration are dorsally tan brown and ventrally cream to light yellow; small black dotting is evident throughout the larva's body; formerly, R. cascadae, R. kezeri, R. variegatus and along with R. olympicus were considered one species listed as R. olympicus; currently, there are no recognized subspecies.

Diet: Consists of small aquatic & semi-aquatic invertebrates and their larvae, such as worms, snails and insects in particular midges and their larvae, other swarming & aquatic larval dipterans, ephemeropterans mayflies) and trichopterans (caddisflies) that frequent waters; the predaceous Dicamptodon tenebrosus shares the same streamside range as the olympic salamander and most probably in all its phases preys on the much smaller salamander's eggs, larvae & adults; garter snakes are also most probably a predator of this salamander.

Habits: This salamander spends most of its time in water or no more than 1 meter/3.3 feet from the water's edge; apparently the olympic salamander are unable to disperse through dry forest, although during rains have been found up to 122m/402.6ft from water source; it is stenothermic, so therefore is very sensitive to heat & desiccation and avoids any direct sunlight; thermal tolerance for Rhyacotriton species is among the lowest for amphibians; fatal critical thermal maximum in water is reached for adults at 27.8 °C/82 °F and even lower for larvae at 26.5 °C/80 °F; during droughts they will seek subsurface flowing water; main travel is more upstream than down; when on the water's edge the white flecking (guanophores) serve as camouflage blending into its background appearing as the glistening dewdrops from overspray landing on the ground cover it is standing on; when threatened or startled exhibits the unken reflex by rolling over and displaying its bright venter; females are found year round internally

3. Guanophore camouflage

containing a male's spermatophore suggesting year round breeding and that oviposition occurs throughout the year with a peak period in May; an individual female does not oviposit fertile eggs throughout the year, but once annually; courtship is in conducted in the water and eggs are laid in clear, cold flowing streams under submerged rock beds, gravel bottoms or narrow rock cracks; on average 8 eggs are laid and no adult attends them; eggs hatch anywhere between 210 to 295 days; the larvae sustain nourishment from their yolk sac for another 295 to 360 days; whereupon after that period need to actively feed; this is the longest embryonic period for any oviparous salamander. larval duration is ~ 3.5yrs with metamorphisis into the adult occurring when the larva has reach 33 to 44mm/1.3-1.7in SVL; the juveniles become sexually active after another 1-1.5 yrs; in their biome, being dependent on a stream microhabitat and as a specialist as opposed to a generalist in livelihood within the microhabitat, any environmental change adversely affects this salamander first before any other fauna; logging in the salamander's range has extirpated this salamander from the logged area due to stream siltation & heavy sedimentation as resulted from the practice of logging; global warming and its affects on climate will immediately have negative impact on this salamander; total population is in decline.

Vulnerable (3.1) IUCN

1. Semirechensk salamander

Semirechensk Salamander
Family: Hynobiidae
Genus: Ranodon
Species: Ranodon sibiricus

Length: 15-25cm/6-10in total length (TL)
Weight: 28.35-141.75g/1-5oz
Longevity: 15-20yrs
Name Origin: Ranodon sibiricus ~ Ran is derived from the Greek word, rana meaning 'frog' while odon is Greek for 'teeth'; sibiricus is Latin in masculine singular form meaning, 'a native of Siberia'; all is in reference to, 'frog tooth salamander of Siberia'.

Distribution: Where distribution once extended well into western Siberian plains, current range is restricted to fragmented populations of the Ala Tau & Tien Shan Mountains of western China & eastern Kazakhstan in plateau brooks, mountain streams, marshes and terrain 1,500-2,750m/5,000-9,000ft asl; prefers flat plateaus dense with permanent brooks & streams.

2. Typical habitat terrain

Description: With prominent labial folds, in my opinion is one of the oddest looking of salamanders; with a lineage dating back to 300 mya, is a true living fossil relic; in studying larval development, information has been invaluable in understanding limb evolvement of tetrapods; developed cartilaginous neurocranium connected with upper part of mandibular arch; is a larger robust salamander exhibiting a prefontal fontanel (deep connective tissue at anterior of skull) and well developed paratoid glands; possesses vomerine teeth set in two short & widely separated series; eyes are big and protruding; lungs are present, but reduced; 11-13 costal grooves are present; there is a dorsal longitudinal groove extending from occipital bone region to base of tail; tail base is cylindrical with remaining portion highly finned tapering towards tip in sword-shaped fashion; males have a dimorphic longer tail and during breeding season, male tail fins exhibit fewer undulations and are higher than females; year round, male limbs & head are more muscular; tail is slightly larger than body in both sexes; no extra protective keratinization on palm & sole coverings; overall dorsum coloration is a yellowish brown to a darker greenish olive gray, while some individuals exhibit a more mottled & spotted pattern; color may change dependent upon temperature and environment where aquatic individuals become darker than terrestrial ones and higher temperatures lighten coloration; close-up the salamander may appear as speckled with glitter; ventrum is always lighter; there are no recognized subspecies.

3. Note nasolabial folds

4. Another close-up

Diet: Semirechensk salamanders are carnivorous eating almost exclusively invertebrates and their larvae; adults consume aquatic & terrestrial insects, while larvae consume primarily aquatic invertebrate larvae; during ontogeny (development to maturity), feeding is divided into 3 stages: endogenous (organism sustenance from self), mixed & exogenous (sustenance from external sources); for endogenous behavior, larvae feed off yolk for first 4 days; exogenous is further subdivided into: consumption of hydrobionts (varying aquatic organisms) by larvae up to full metamorphosis, consumption of terrestrial invertebrates by recently fully metamorphosed juveniles and consumption of terrestrial & aquatic invertebrates by adults; R. sibiricus is considered rheophilic (lives in aquatic environments with current & flow), but larval diet consists of numerous limnophilous (living in still waters as lakes & marshes) organisms; most likely is due to salamanders's preference for slow currents in brooks that attracts limnophilous invertebrates; the terrain inhabited protects this salamander from predation, as the land is too rugged and cold to support predators; about the only predation are adults & larger larvae cannibalizing the smaller larvae.

Habits: Sub-adult & adults are mostly nocturnal while larval stages spend a lot of active time diurnally; despite being rheophilic, the robustness, instead of more streamlined body form, R. sibiricus resembles pond salamanders more than salamanders living in rushing waters; while terrestrial adults hide under loose grasses in wet meadows or beside streams, in holes or under loose stones, larvae hide under submerged debris or rocks; hibernation for adults is on land underneath sheltering rocks or shallow holes; since achieving full metamorphism may take up to 3 yrs, depending on altitude & climate, larvae must hibernate as well underwater; terrestrial individuals begin hibernation in late September to early October remaining in a torpid state until spring comes; from April to June, adults come out of hibernation as soon as snow & ice begins melting migrating to their ancestral breeding water sources; spawning takes place from

5. Keep on trucking lil' fella

April to August; males first arrive into the water waiting for females, whereupon once females enter water, olfactory senses alert males and leads him to flanks & tail of female where he gently bites her; when egg deposition nears, female then approaches male; it is the male in charge of finding and choosing a suitable nesting site; once site is established, males deposit spermatophores on stems of submerged plants or undersides of rocks in size from pebbles to boulders; even though spermatophores are designed for internal fertilization, fertilization is external with female depositing her egg sacs with the aid of a mucoid stalk alongside the spermatophores, thus fertilizing the eggs as spermatozoa begins floating & saturating area; female produces 2 egg sacs each containing ~ 32-58 opalescent, bluish translucent eggs; upon hatching; the larvae are 17-20mm/0.67-0.79in; latter stage larvae with limbs possess chitinous claws, but lose them after final metamorphosis; deforestation, livestock trampling eggs, larvae & adults, human consumption for worthless medicinal purposes and over harvesting for scientific studies & the pet trade have made this salamander's fragmented population highly threatened; its already small range is declining still; this salamander's viability is in the worst condition of all salamanders from the former Soviet Union with populations of China critically endangered.

Endangered (3.1) IUCN

Frogs:
Frogs and toads which are specialized frogs, all are under the order, Anura. The distinction between what is a frog and what is a toad is purely based on appearance rather than taxonomic value. What frogs we label as toads are primarily in the family, Bufonidae described generally in possessing dry warty skin and with short legs used more for hopping than jumping. Adaptations to drier environments by toads is an example of convergent evolution in adaption to a new ecological biome. Also, where most frogs don't, most toads possess a pair of cranial crests that are bony ridges on the dorsal side of the head and bulbous parotoid glands (alternatively written as paratoid), which are an external gland that secretes a neurotoxic alkaloid known as bufotoxin.

1. Agalychnis callidryas

Among other traits, with most equipped with a single vocal sac, frogs are probably best known for their vocalization. But frogs don't just rib-it, they also croak, chirp, peep, scream, whistle, bark, howl, hoot and quack while some species sound like an angry cat with Nyctibatrachus poocha even meowing like a cat.

Sounds emitted by frogs are articulated by various species for a specific form of communication. In calling to females, males are advertising in saying I'm well and I'm here for you. Although these are the frog calls most of us are familiar with, anurans do indeed communicate in different calls for other reasons. There are aggressive male calls towards another male, distress calls when first captured by a predator and male release calls to other males in saying, "Hey, I'm a male too, get off my back." For you see, during mating time, things can get a little crowded and hectic in a pond full of enamored frogs. In certain species, females emit calls as well.

Frog lineage goes all the way back to the Permian 275 mya. As explained in the salamander discussion section, Gerobatrachus hottoni had both salamander and frog characteristics. The first proto-type frog occurred, so far in the fossil record, during the early Triassic around 250 mya in what is now Madagascar. This animal, other than having a short tail and unable to yet

2. Triadobatrachus massinoti

jump, had many other frog characteristics. Known as Triadobatrachus massinoti, the tibia and fibula were beginning to elongate for jumping capabilities. As in true frogs, it had an anteriorly directed ilium, a reduction in number of presacral vertebrae, the fusion of the parietal and frontal into one bone and was toothless in the lower jaw.

The earliest known 'modern frog' dates back to the Lower Cretaceous 125 mya. Found in the western part of Liaoning province of China from the rich fossil beds of Sihetun was an animal known as Sanyanlichan, which may be a direct ancestor to the frog family, Discoglossidae. Except for having one extra presacral vertebrate, for all intent, it appeared as a modern day frog, but due to a few salamander morphological traits, it did not reach the status of 'fully modern frog.' But after the appearance of Sanyalichan, a whole slew of truly modern frogs began to appear.

3. Prosalirus bitis fossil

The extinct genus, Vieraella produced some of the first 'modern frogs' actually first showing up around 200 mya during the Lower Jurassic. Though not the direct descendant of today's anuran frogs, it was a fully formed frog. An example of this genus is the extinct, Vireaella herbsti, which was very small from snout-to-vent at only 3.3cm/1.3in.

Prosalirus bitis had a skeletal structure designed to absorb the force of jumping with long hind limbs. Also from the Lower Jurassic, it lived 190 mya and is tentatively placed as the direct descendant of anurans.

As previously stated, once the genus, Sanyanlichan arrived 'fully modern frogs' exploded onto the scene. Two of the earliest 'fully modern frogs' taxa were Callobatrachus and Mesophyrne from the Jurassic/Cretaceous border 144 mya, followed by Mid-Late Cretaceous (120-65 mya) examples by the likes of Shomronella, Saltenia and Estesius and Tertiary forms by way of Limonodynastes and the families of Paleobatidae and Ranidae 65-2 mya.

All this frog speciation lead to the largest known frog, Beezlebufo ampinga, or 'devil frog.' Weighing in at 4.5kg/10lbs and reaching a length of 40cm/16in, this frog was essentially all mouth and one big stomach. Living some 70 mya, it would lie in wait and ambush pretty much any small animal that happened by. It is for sure that its menu also consisted of baby dinosaurs or other little theropods.

4. B. ampinga eating a dinosaur

Except for the extant species in the primitive family, Ascaphidae, today's adult frogs lack a tail. With large hind legs for hopping or jumping, their forelimbs are also well developed and aid in tetrapodal walking. There is no nuchal region (neck) with the head appearing as being directly attached to the body.

Though many of the young or tadpoles are omnivorous or even herbivorous, all adults are carnivorous. Prey has to first move, for all frogs only recognize motion as a potential meal and will eat anything that moves and can be swallowed. Distasteful, acrid or foul smelling animals, such as salamanders and certain insects, once captured in the mouth, the frog will immediately spit and/or claw with the forelimbs the once intended prey back out.

Since true frog anatomy came onto the scene, there was an explosion of speciation that radiated into all climates, elevations and terrain. From tropical to subzero, there you will find frog species. Most reside in an elevation range of from sea level to 2,000m/6,600ft, with many reaching up to 4,000m/13,200ft. But Telmatobius, a genus found in the Peruvian Andes, has species that surpass 5,000m/16,500ft.

Most species are relegated to tropical environs due to water dependency, but the hottest of deserts have been conquered as well, with species that have evolved deterrents to desiccation. This is a huge milestone in achievement to have been successful in coming from watery descendancy to surviving in regions that receive less than 7.5cm/3in per year.

As most of us are taught, frogs in reproducing gather around water bodies every spring to perform amplexus where the female and male releases her eggs and his spermatozoa simultaneously in water fertilizing the eggs, where they will soon hatch into tadpoles possessing gills. The tadpoles go through various stages of aquatic metamorphism until finally reaching the lung air breathing adult stage that may now live on land.

Due to natural selection in specific environments, other reproductive strategies have evolved that perhaps we are not too familiar with. The tree frog family, Hylidae has members that do not mate en masse in larger and more permanent bodies of water, but rather in moist forests lay externally fertilized eggs in pools of water collected in broad leaves such as bromeliads. Though there could be risks in the puddle drying up or the leaf developing a tear and thus leak, this strategy greatly reduces competition and predation.

Other hylids, such as Oophaga pumilio will lay their externally fertilized eggs on land and after hatching show some parental care by carefully taking the tadpoles to water pools and tend them. Still, there are those that lay eggs both on water or land. Dendropsophus ebraccatus, otherwise known as the pantless treefrog because a part of the hind limbs lack pigmentation, is transitional in laying eggs in either aquatic environs or terrestrial ones. The eggs are not designed to excel in either environment. The risk on land with these adapted eggs is that they may desiccate, but the risk in water is not enough oxygen uptake. Though it has not been studied, I propose the female adults most likely will lay eggs on land when moist conditions prevail, but in waters when water temperatures are cooler retaining more dissolved oxygen.

5. D. abraccatus aquatic egg laying

6. D. abraccatus land eggs

Other tree frogs lay eggs on overhanging branches or large grass stems over water whereupon hatching, the tadpoles drop into the water below, while others will lay eggs in foamy wt nest on leaves or above water. Furthermore, there are species that lay eggs in self dug moist holes or in already made dips where rainwater run-off will take the eggs to water. In some of these land egg laying species, the eggs fully develop into miniature froglets.

There are frogs that lay eggs onto there bodies in pouches, onto the male's hind legs or simply adhered to the back, or even placed inside the body. In the Surinam toad (Pipa pipa) individual eggs are placed in pits on the female's back where once hatching the tadpoles remain until fully developed. Males of Rhinoderma darwinii brood developing young in their vocal sac until fully metamorphosed. With two species in the genus, Rhehbatrachus, the gastric brooding frog of Australia, which is feared to have just become extinct, incubates prejuvenile stages of offspring in the stomachs of the female where she shuts down all ingestion and digestion of food during this maternity period. Once the froglets are fully developed, the female regurgitates them out into the world.

Yet, there are still unique frogs that have developed ovoviviparity and even viviparity. For an ovoviviparous example, females in very rare species of the genus, Eleutherodactylus retain their internally fertilized eggs in a distended section of the reproductive tract. There the eggs hatch and the tadpoles fully metamorphose until the miniature froglets are born. In the Tanzanian toad genus, Nectophrynoides females actually have a nourishing yolk placenta that feeds developing fetuses that arose from internally fertilized eggs. These toads give live births to miniature toads.

Adult frogs depend heavily on the senses of sight and sound. In hearing, frogs do not possess external ears, but do have a well developed external tympanum located just behind and below each eye. As a thin membrane surrounded by a cartilage ring, as sound hits the tympanum it vibrates correspondingly. This in turn relays the produced vibrations to a rod transferring the sound waves to the inner tympanum. Inner ear fluids are disturbed washing waves over onto cilia (tiny hairs) that are connected to nerve fibers. These nerves convert the sound wave information as electrical impulses that are transferred to the auditory portion of the brain for interpretation.

7. Rana clamitans tympanum

Frogs also hear with their lungs. As sound waves hit the lungs they're immediately directed to then tympanum via an air-space link. This method allows the frog to determine sound location from the difference in pressure of external environment sound against the same sound being funneled through the lungs. The air link between the lungs and tympanum also serves as a buffer by equalizing pressure differentials from the internal and external portions of the tympanum which keeps it from rupturing. Frogs themselves can emit croaks up to 95 decibels which is as loud as a jackhammer.

The bulging eyes of a frog allow it to see almost simultaneously in all directions in allowing it to assess it's environment in the detection of prey and in the avoidance of predators. Without a neck to turn the head, the functionally directional bulging eyes are a frogs main asset in staying in tune to its surroundings. Relying heavily on sight, the pupil picks up enough light to allow for night vision and there is a thin clear membrane that involuntarily closes over the eye when its in water serving as protection. The frog also has a functioning eyelid that may be voluntarily closed over the eye at anytime the frog desires. The eyes are not equipped for long distance vision, but the directional eyesight does not overlap, so a nearby flying insect is readily assessed and snagged right out of mid-air.

One other function of the bulbous eyes are to aid the frog in swallowing its meals. Once prey is in the process of being swallowed by the frog, the eyes, appearing as blinking in closing the lower and upper eyelids together, sink down deep into the socket. Since the frog has no teeth for chewing nor a functional tongue for pushing food down its gullet, the recessed eye balls aid the animal in swallowing food by applying pressure downwards into the throat. Though there are a few exceptions and even one family (Pipidae) with four genera that are tongueless, for a catapult effect, a typical frog tongue is attached in front while free in the back.

Frogs in temperate climes will hibernate during the colder months usually buried in mud bottoms respiring through their skin. Rana sylvatica, whose range extends past the Arctic Circle, has the ability to survive freezing. In more arid conditions frogs may aestivate for months burrowed in desert soils during the hottest months and not appear onto the surface until the next rains come.

Under sixteen families, there are more species per genus members than found in any other animal group. For instance, under the genus, Scinax there are currently 106 species, while in the genus, Litoria there are 160 species. The genus, Eleutherodactylus though, has over 500 species members.

1. Black Toad ~ Bufo exsul

Black Toad
Family: Bufonidae
Genus: Bufo
Species: Bufo exsul

Length: ♂ ~ 44-59mm/1.7-2.3in SVL; ♀ ~ 46-69mm/1.8-2.7in SVL (snout-vent length)
Weight: Mean ~ 13.75-25.56g/0.49-0.90oz
Longevity: Maximum 11.5 yrs
Name Origin: Bufo exsul ~ Bufo is Latin for 'toad'; exsul is Latin for 'exile'; altogether means 'exiled toad' in reference to the species' isolation.

Distribution: Endemic to 4 spring systems all within Deep Springs Valley in Inyo County, California; the springs in the valley are: Bog mound, Buckhorn, Corral & Antelope Spring; attempts have been made to introduce populations to other comparable spring regions, all have failed except one with a managing small population in a flowing well near Salt Lake in Death Valley National Park; natural range is one of the smallest for all N. American amphibians covering only 150,000sq meter/15ha; prefers natural springs or seepage in arid regions straying no more than 12m/39.6 ft from water source; will not frequent water portions that are rocky or provide no shade; normally in or near waters shaded by juniper, pinon pine and tussock.

2. B. exsul ventrum

Description: Since 2006, there is a push to change genus name from Bufo to Anaxyrus; is a close relative to Bufo boreas; is a smaller toad and more water loving than most toads; skin is dry and though with warts is relatively smooth for toad standards; does not possess cranial crests (bony ridges on the heads of toads); parotoid glands are oval in shape; Overall coloration is black with light tan or white spotting. Some individuals may be virtually solid black on the dorsal half; There is less black on the ventral side; throat region usually has the least black displaying only dark specking over a palish white; a light stripe running down the center of the back is normally visible; juveniles are not dark, but olive green instead. There is no recognized subspecies.

3. Virtually black individual

Diet: Consumes a variety of small invertebrates primarily insects including beetles, ants, bees, flies and the larvae of dipterans (flies), lepidopterans (butterfly caterpillars) & coleopteran (beetle) grubs; recognize prey through sight then pounce and capture prey with sticky tongue; spiders, mites, collembolla (springtails), hymenopterans (wasps), fairy shrimp and homopterans (cicadas) when available; tadpoles consume organic & inorganic material found floating or suspended in water, attached to plants or laying on muddy bottoms, but may capture a small enough invertebrate that it can pass through its mouth and gullet; there are no known predators, but eviscerated black toad corpses have been observed with fatal crow pecking marks.

Habits: Living in very arid confines that can bear witness to extreme cold and hot conditions, this toad is active from March to September; spends winter underground in rodent burrows near water source in torpor hibernation. may overwinter with many individuals in same burrow; is primarily diurnal, but during hotter months becomes crespuscular while during hottest months is nocturnal; does not aestivate; for good reason surrounded by arid lands with spring pools acting as an oasis, does not venture far from water's edge; is a viable swimmer swimming underwater more like frogs than toads; will refrain from entering stony rock bottoms, preferring instead to remain in aquatic areas of common black muds where its body coloration blends in

4. B. exsul preferred habitat

well; prefers walking or slowly crawling as opposed to hopping; forages along water edges, but will venture into nearby tussock growths for food; has no vocal sac, but males are still able to emit weak high-pitched chirps to address territoriality, in particular during breeding season; breeding extends from a period of late March to early May, breeding is aquatic conducted in the shallows no more than 20cm/7.87 in in depth; during amplexus fertilization is external; eggs are laid out in strings and intermingling with aquatic plant stems; eggs hatch into tadpoles after ~ 5 days of being laid; length of larval stage (tadpole) is from 3-5 weeks; tadpoles do not hibernate

5. Juvenile coloration

overwinter, but fully metamorphose into juveniles reaching 22mm/0.87in before winter months arrive; once developed, juveniles remain alongside adults in marshy areas of anaerobic black mud; when alarmed will immediately swim to water bottoms where they camouflage well with the mud, if caught from water's edge will attempt to find and enter first hole available among the tussock growth; for extremes in climate, appear to have adapted well; though populations seem stabilized, due to a very restricted & isolated habitat that is specialized, this toad is considered as threatened; if the small springs ever dried up or a viable predator was introduced, this toad would become extinct; cattle occasionally come to drink at the springs or graze the tussock bunch grasses, but properly managed most probably is beneficial as their manure attracts potential insect prey for the toads.

Vulnerable (3.1) IUCN

1. Mexican burrowing toad ~ R. dorsalis

Mexican Burrowing Toad
Family: Rhinophrynidae
Genus: Rhinophrynus
Species: Rhinophrynus dorsalis

Length: 5.1-7cm/2-2.75in; record: 8.9cm/3.50in
Weight: 2.5-4.0g/.09-.14oz
Longevity: In wilds unknown; a captive at Fort Worth Zoo lived 2yrs & 2mths
Name origin: Rhynophrynus dorsalis ~ Rhinos is Greek for 'nose' in reference to peculiar snout & phrynos is Greek for 'toad'; dorsalis is Latin referring to 'of the back'; altogether refers to odd appearance.

Distribution: Central america is the core range extending from the southern end into Costa Rica all the way up through the Gulf Coast eastern corridor of Mexico into the extreme southwestern Texas lowlands of Starr & Zapata Counties and up halfway along Mexico's western Pacific corridor; prefers dry or arid friable soils for its fossorial lifestyle as opposed to forested areas. found in coastal plains to elevations of 500m/1,650ft.

2. Close-up of odd face

Description: This primeval amphibian is no more a modern toad than a frog is a salamander; humans are genetically tied closer to rodents and bats than R. dorsalis is to modern day frogs and toads; based on evolutionary outcrop groupings, the ancestor to all extinct and modern anurans could have had a larval stage that greatly compared to that of a rhynophrynid tadpole; with 190 million years of evolutionary engineering, this amphibian has retained morphological visages of extinct ancestors from long ago; snout is covered with unique epidermal spiking; buccal & esophageal linings are ornately folded; lips act as a double closure along the long wedged shaped edentate (like edentate mammals with few teeth such as armadillos & anteaters) maxillary arch; the mechanism for tongue protrusion is distinct from other frogs that project their tongues from lingual flipping; with rhynophrynids,

3. Juvenile coloration

intrinsic tongue muscles stiffen the organ by exerting hydrostatic pressure on the fluid contents of lingual sinus sending sticky tongue outwards in rod-like and edentate fashion; the tongue is rooted in the back and not in the front as most frog tongues are; there is no apparent external tympanum and eyes are tiny; head is small with cone shaped snout; body is stocky and inflatable held up by short & fat limbs; forefeet are not webbed while hind feet are heavily webbed; overall coloration is a dark purplish-gray with scattered orange spotting and a mid dorsal orange line running down its back; juveniles are more gray with no orange spotting and displaying a lighter gray mid line that has yet no orange; this species is not only in a monotypic genus, but its family is monotypic with no recognized subspecies.

Diet: Appears to exclusively feed on subterranean ants and termites as its mouth, tongue and gullet display adaptive specializations in feeding on small insects; actual protrusion of edentate-like tongue through buccal groove to snatch prey, involves shifting the tongue forward via protraction of hyoid by extrinsic muscles to the tongue; tongue protrusion maechanism is uniquely designed for underground advancement on small insect prey items; front limbs & spiked snout are also specialized for entering termite/ant burrows & chambers; tadpoles are phytoplankton feeders; with their strict fossorial existence there aren't any known predators that prey on adults, but when surfaced, it is quite possible large enough snakes would consume even inflated adults; eggs and tadpole predators range from amphibians, reptiles to other temporary pool inhabitants including other tadpoles; R. dorsalis tadpoles may resort to cannibalism if there is no food to be had, with larger tadpole groups attacking smaller ones.

Habits: This species is strictly fossorial only surfacing during rains or after heavy rains to breed in temporary ephemeral pools; when alarmed has ability to increase body size by 50% by

4. Anterior view of inflated body

inflating lungs with air; this deters predators in making the body appear larger and is more rigid; another defense is a sticky & foul skin secretion that also serves as a deterrent from marauding ants & termites when the frog is attacking their domiciles; consumes termites and ants much like an anteater mammal would with a protruding rod-like tongue being ejected out from the mouth via the buccal groove (as discussed above) that serves as a vaulting point located in the roof of the mouth; the tongue exits and enters the mouth through a small groove located at the front of the lower jaw; this frog burrows backwards using his hind limbs which are equipped with keratinized tubercules for digging burrows; as the frog continues burrowing, the body is very flexible and can maneuver easily or shift around rock or hardened dirt clods; when digging, body is twisted and change in inflation patterns are employed to facilitate penetration of soils; in doing so, the filling of the shaft from where they were in burrowing is filled, allowing no entry for potential predators; when approaching a potential prey area, turns around and enters foreward with spiked snout and front limbs doing the digging; with snout in position of insect tunnel and when prey are detected, the normally flat irregular tongue is reshaped into rod fashion by the muscle previously discussed and then

5. Dorsal view of inflation

extended outwards where prey are captured on cup-shaped tip of tongue and drawn back into mouth; action by the sterno-hyoid pulls the tongue back in; grooves in roof of mouth and the esophageal folds serve as protection from mandibular prey bites by disorienting the insect prey; further, the esophageal walls are lined with cells that insulate more vulnerable portions and are rapidly replaced if damaged; the only reason this frog can eject its tongue in such a matter like certain salamanders, chameleons and other lesser publicized tongue feeding lepidosaurs is strictly due to convergent evolution and not genetic relationships as the tongue protruding mechanisms are different; possessing torpor stability, it can go for extended periods of not eating, where in excess of 18 mths. have been observed; also can survive dehydration in an aestivative state and will fully recover once rehydrated; typical of nocturnal foragers, the eye pupils are vertically slit; does not attempt to make cocoons as some other species of burrowing toads do during aestivation of hotter months; appears not to even aestivate or hibernate as can be dug up throughout the year as active; during the wet season may be found just underground at 7-15cm/2.76-5.91in; self made chambers are found a bit deeper underground during drier periods; breeding may occur at any time of year followed by heavy rains filling in temporary bodies of water of ephemeral pools, ditches, culverts and flooded fields; males may begin their mating calls while still burrowed underground; the male call is a whooaaa much like the

6. Note: mouth barbels

ol' cowpoke telling his horse to stop; once above ground, many migrate up to 1.6km/1.0mi in mass to the seasonal waters; males intensify their mating calls near water's edge or while floating; amplexus soon ensues and with external fertilization, up to 8,000 eggs are laid in sticky clumps that sinks to bottom, but soon segregate and individually rise to surface; within 5 days eggs hatch with tadpoles emerging toothless and no mouthparts; tadpoles are filter feeders with barbels directing tiny diatoms and algae into oral cavity; tadpoles form large schools with mass aggregates of varying sizes or stages of development; tadpoles form large shoals simultaneously facing the same direction; in doing so, their synchronized undulating movements creates currents stirring detritus material from bottoms putting food within reach of their barbels for ingestion; tadpoles in much smaller schools do not grow as quick and remain smaller than their counterparts in the larger schools; schools may contain 1,000s of individuals and are disc-like in shape reaching a depth of 25cm/9.84in; once tadpoles metamorphose into a juvenile adult form, they may still retain their tail but immediately leave the water to burrow hiking the tail up on the back to allow the hind legs to dig without interference; due to fossorial lifestyle, there is no population estimate, but numbers are believed stable making listing as least concern.

1. Red-eyed treefrog ~ A. callidryas

Red-eyed Treefrog
Family: Hylidae
Genus: Agalychnis
Species: Agalychnis callidryas

Length: ♂~ 51-64mm/2.01-2.52in ♀~ 64-77mm/2.52-3.03in
Weight: 6-15g/0.21-0.53oz
Longevity: Observed 4yrs & 1mth in captivity but in wilds is suspected longer at 8-10yrs
Name Origin: Agalychnis callidryas ~ Agalychnis is derived from 2 Greek words, aga meaning 'plenty of' and lynchos meaning 'to shine' in reference to bright eyes; callidryas is derived from 2 Greek words, kallos meaning 'beautiful' and dryas meaning 'tree nymph'; altogether refers to, 'beautiful shining-eyed tree frog'.

Distribution: Most dense populations are massed on the Caribbean side of Central America but extends from Mexico's Yucatan Peninsula through Central America down to its southern most range in extreme northwest Colombia; prefers tropical lowlands with humid forests near rivers and permanent streams.

Description: Most astounding feature to this medium sized tree frog is its bulging red eyes

2. Asleep; note eyelids & toepads

3. Golden reticulated membrane

with vertically slit dark pupils; 3 reticulated membranes (eyelids) on each eye appear darkish purple with numerous random light golden webbing; have long spindly legs in comparison to thin streamlined body; skin is smooth with a rounded head and truncated snout; adhesive toepads exist to accommodate holding on when climbing; mouth is oral disc in shape with serrated beaks; there are 2 upper & 3 lower rows of denticles; although A. callidryas, belonging to phyllomedusine (leaf breeding frogs) do not possess highly toxic compounds, they do contain high levels of biologically active peptides; as coloration varies in populations from one locality to another; dorsum and outside legs are normally bright lime green with inside of legs blue; dorsal side can also be predominantly a darker green with tiny whitish cream spotting; on the sides are blue to deep purple background interrupted by vertical yellow, cream or white striping; feet are orange or deep red in color; with only population variants, juveniles are brownish to green; there are no recognized subspecies.

Diet: Are carnivorous and ambush predators eating crickets, flying insects and any other small insects that frequent tree canopies; have been known to take other smaller tree frog species; tadpoles eat tiny aquatic animals such as crustaceans, fruit flies and pinhead crickets; from carnivorous birds to snakes, many predators prey on these nonpoisonous tree frogs; arboreal snakes such as Leptophis ahaetulla & Leptodeira septentrionalis specialize in preying on whole egg clutches; the social wasp, Polybia rejecta will attack one egg by plucking the embryo, but in numbers can decimate an egg clutch; tadpoles coming from eggs laid on overhanging plants are exposed to many aquatic predators when they drop into the water; eggs also succumb to fungal diseases, while tadpoles can die from stagnant puddles due to lack of dissolved oxygen (DO); many newly metamorphosed froglets are taken by water spiders hunting on water's surface or edge.

Habits: Is strictly an arboreal nocturnal frog active at night and sleeping by day in the tree canopies; as how legs are designed, they prefer to walk as opposed to jumping through canopies; toe pads are designed to cling to smallest twigs or largest trunks; to avoid detection

4. Darker green specimen

when asleep during daylight, the frog picks a green background to rest on, tucks its hind legs along and over its sides concealing the bright lateral body coloration, folds in and lays over its bright feet and in closing its bright red eyes with only its green back exposed, blends in well, thus being concealed and inconspicuous to its surrounding environment; if alarmed though by a predator, it will instantly and simultaneously open its red eyes and expose its colored body parts; this startling mechanism effect may surprise the predator just enough, for the frog to make for a quick get away; mostly, the frogs sleep underneath broad leafed plants; virtually the only time red-eyed frogs descend from the tree canopies is during mating; the breeding season occurs

5. Close-up of face

between from late May to November coinciding with the wet season; equipped with 2 vocal sacs, males are the first to descend and begin their chuckling aggressive sounds of, 'chak, chak-chak' to setup territories and advertisement calls to attract mates; preferring quiet pools with plenty of overhanging vegetation, the males set up territories from edges of ponds and slow backwaters or low lying plants during wetter nights, while most may be perched higher up during drier times; when competing, males display an act to communicate with one another known as tremulation; this aggressive signaling from one male to another is conducted by creating vibrations within the substrate it is on, which is usually a leaf, or by shaking a small twig rapidly with hind legs; communicative tremulations occur when 2 males are within 2 meters/6.6ft of one another and only occurs when both are on the same plant; when tremulating, it will continue in reciprocal fashion with each male signaling, but when there is visual contact during the sending of vibrations, physical wrestling is pursued; once precipitation and water levels are adequate most males head for the waters to lead the females; females descend the canopies very deliberate with one forefoot over the other; on occasion, both sexes have been observed parachuting down to the surface; females approach selected calling males in a straight line and is first come first serve; but there is plenty of activity to go round; once amplexus occurs, the larger female will carry the male into the water to absorb water through her skin and into her bladder as an ingredient in making the jelly mass surrounding her eggs; once this occurs, the female still in an amplectant embrace with male then moves up into the overhanging plants in search of a suitable site for egg deposition;

6. Amplectant pair sleeping

the female looks primarily at the undersides of broad leafs or other plant parts such as vines that overhang above the water; unusual sites have been observed as well like in water-filled larger animal tracks, even human footprints; egg clutch sizes range from 20-50 green eggs with a mean average of 40; the female may lay up to 5 clutches per night for up to 3 nights; the male will remain in amplexus with the female throughout the term even while sleeping and fending off other males who try to take over when the female returns to water to rehydrate; the eggs are immediately fertilized externally by the male as the female oviposits; at times eggs are laid on topside of leaf where the apparently the leaf curls in over the egg clutch on its own attached by the adhesion of the jelly mass; most leaf breeding tree frogs intentionally curl leaves over the egg clutch for protection in concealment and dehydration, but for the red-eyed tree frog, it appears the frog is aided by the plant itself; when laid over water up to 6.06m/20ft high, embryo development occurs synchronously but hatching occurs asynchronously in 6-10 days; when left undisturbed the eggs hatch into tadpoles on a mean average of 6.5 days; many disturbances can occur though that affect and accelerate hatching down to four days; snakes that will consume the entire clutch are detected by the developing tadpoles through specific vibrations the snake makes causing the entire clutch to hatch with the tadpoles escaping by falling into the water; the social wasp only attacks one egg at a time; the intended tadpole in the egg sought after will again detect the wasp's specific vibrations, hatch and fall into the water while the rest of the embryos remain in their eggs; a developing tadpole can even detect the minute vibrations sent out by the developing spores of fungus infestation by a filamentous ascomycete (Phaeosphaeriaceae); in egg clutches not affected, random hatching occurs; in those that are infected, nonrandom

7. Egg clutch

hatching occurs; eggs in direct contact with the fungal hyphae hatch and with the empty egg casing, essentially form a barrier from further invasion of infection into remaining unhatched eggs; in every instance the embryos do not instantaneously hatch once vibrations are detected, but in a sense evaluate what is occurring before risking premature hatching; some tadpoles hatch then drop to the ground instead of water; the larvae (tadpole) can live up to 20 hours out of water, and usually since hatching occurs during wet season, rains wash the stranded larva into the water pool source; tadpoles show rapid external gill loss and may lose their outside gills within 3 minutes upon hatching, relying on less functional internal gills to breathe with; even so, the advantage to this is that external gills create drag, so without outside gill structures, tadpoles can now swim more rapidly and efficiently; tadpoles suspend themselves vertically in water column to filter feed; once tadpoles have fully metamorphosed in 75-80 days, rather than red, the froglets possess yellow eyes and change body color in being green by day to purple or reddish brown by night; though habitat destruction has had its impact on regional populations and has been heavily sought after in the pet trade, there are no current threats to its continued existence.

1. Golden dart frog ~ P. terribilis

Golden Dart Frog

Family: Dendrobatidae

Genus: Phyllobates

Species: Phyllobates terribilis

Length: Ave snout-vent length (SVL): ♂~4.5cm/1.77in; ♀~4.7cm/1.85in; max: 5.5cm/2.17in
Weight: < 28gm/1oz

Longevity: > 10yrs

Name Origin: Phyllobates terribilis ~ Phyllobates is Greek as 2 words conjoined meaning phyllos: 'leaf' and bates as 'climber' in reference to an arboreal lifestyle; terribilis is Latin as 2 words conjoined meaning terri: 'terrible' and bili as 'anger' referring to aggressive nature; altogether means, 'frightening tree climber'.

Distribution: Is endemic to Columbia on a Pacific coastal terrain in the western foothills of the Andes' northerly inclined spur of the Cordillera Occidental; are found in elevation from 90-200m/ 297-660ft throughout rainforests from very moist slopes to drier ridge tops; preferred terrain is rough very steep slopes that are sometimes even with perpendicular inclines of 90 degrees; is always found near rivers, but now mostly near smaller streams as agricultural clearings near larger rivers have decimated required habitat.

2. Mint green morph

Description: Though a small frog, is the largest of dendrobatids; snout is sloping and rounded in a lateral profile and bluntly truncated dorsally; unique to Phyllobates genus species, possesses a bony plate in upper jaw that substitutes for teeth; teeth are present on maxillary arch; loreal region is both vertical and slightly concave; there are 4 toes on each forefoot with third toe as largest and 5 on each hind foot; hind toes descend in size order from 4 > 3 > 5 > 2 >1; each toe ends in a tiny adhesive disc for clinging to arboreal substrate; forefeet have a large rounded palm tubercle; in the hind feet, the third distal tarsus displays a continuous ridge from

3. Pair of white mint morphs

the inner metatarsal tubercle to the tarsal tubercle; body skin is smooth to finely granular while becoming rugose (coarsely granular) on upper hind limbs; tympanum partially concealed in its dosolateral position; this aposematic (poisonous coloring advertisement) comes in 3 color morphs: yellow, mint green & orange and are range specific; the most common is yellow found in the region of Columbia's Quebrada Guangui with shades from pale yellow to a deep golden hue; the yellow morph is the basis for the common name; the mint coloration, existing in the La Brea area of Columbia may be from a metallic green, to a very pale green or even white; the more uncommon morph dispersed throughout Columbian populations is the orange ranging in intensity from a pale yellowish orange to metallic orange; there are spotted and lined melanistic patterns displayed in all 3 morphs; the ventrum is normally a lighter shade of the dorsum coloration; though these color morph variants exist, there currently are no subspecies listed.

4. Orange morph

Diet: This aggressive frog will devour anything it can swallow but primarily subsists on small insects consisting of ants, beetles and termites that abound in its habitat; fruit flies, small young crickets and small lepidodopterans may also be consumed; tadpoles are omnivorous feeding on algae, insect larvae & their mother's infertile eggs; due to highly concentrated levels of poison, there are virtually nil predators that intentionally prey on this frog; the 50cm/19.66in small snake, Leimadophis epinephelus is the only known predator to actively pursue and purposely consume P. terribilis; this snake has built up resistance to this frog's toxins and other frog specie toxins, but is by no means totally immune; once ingesting a golden dart, the snake will go into rapid convulsions then remain motionless for awhile; though the snake is dramatically affected, it still nonetheless remains an astronomical feat; only two-tenths of a microgram (smaller than the pointed end of a pin tip) of just one of the poisons known as batrachotoxin is enough to kill a human once entering the bloodstream; on average 1100 micrograms are produced on a frog at any one time.

Habits: P. terribilis is distinguished by being 'one of the most' on many fronts; given its toxic steroidal alkaloid cocktail of batrachotoxin, batrachotoxin A and homobatrachotoxin, it is not only the most toxic dendrobatid, it is the most toxic of all frogs and one of the most toxic animals of the world on land and sea; it is easily the most aggressive phyllobates in its defiance of predators preferring to simply stand its ground or walk or hop away as opposed to hiding; it also easily resorts to aggression with other P. terribilis outside its social network or other frog species that dare to enter its territory; it is the largest poison dart frogs and one of the largest tree frogs; in blocking cell's sodium channels this frog's poisons are highly neurotoxic and cardiotoxic; this frog carries the most concentrated and potent non-peptidal neurotoxin known on Earth; the frog is totally immune to the poisons it carries as its specialized sodium channels are unaffected; curiously there are 3 insectivorous or omnivorous birds, Pitohui spp., Ifrita kowaldi and Colluricincla megaryncha that, acting most probably as a predator deterrent and insecticide, carry batrachotoxin either within and on their feathers and skin; another animal that carries batrachotoxin are certain beetles from the family, Melydridae; all three birds and in particular the frog consume these melydrid beetles and is assumed the major source of the toxin; P. terribilis also consumes myrmecine & formicine ants; except for the feet, all the body's skin contains glands that store the toxin where it is secreted coating the body's skin; the poison is used by the frog in protection & defense only and not for capturing prey; indigenous people such as the Choco & Cofan, by fire, heat the frog causing it to exude small droplets of toxin; once enough poison is collected, arrowheads and darts are then immersed & soaked having a lasting lethal potency of up to 2 years for hunting and warring expeditions; the toxin is long lasting in resisting degradation; even though captives had not been in the wilds for 2 years and had not been fed their dietary toxic source of ants & beetles, they still retained 40% potency; captive bred juveniles and adults, wild tadpoles & newly metamorphosed wild froglets do not possess the toxin; this frog is primarily diurnal and forages as much on the ground as it does in the tree canopies; P. terribilis is one of the most social of all frogs with wild specimens living in groups of 4-7; in being social, they have to interact and have created forms of communication; specific calls, gesturing and touching are all, forms of communicating with one another; push-up movements are a sign of dominance, while head lowering is a sign of submission; calls are more of a trilling sound with both males & females able to call; males however have a shallow subgular vocal sac with small expansion wrinkles at base of throat and well developed paired vocal slits on floor of mouth; trills can be long & sustained and consist of rapid succession of individual notes; frequency can be lowered deeper than other Phyllobates, but can also range upwards to high pitched squeks; breeding in their heavy annual rainfall environment is year round and can breed as often as once a month; family groups converge in mass once or twice a year where heavy aggressive male competition occurs for mates; females remain on the sidelines and fairly calm throughout

5. A male mint with tadpoles

this ordeal during preparation of a male victor; eggs are terrestrially laid where female will expel stored water from her cloaca if there is not an existing puddle; she will also deposit some unfertilized eggs if tadpole's natural food is not abundant in pool; less than 20 eggs in a clutch are laid beneath leaf litter to conceal the site; P. terribilis, as far as frogs are concerned are one of the most devoted parents; during embryonic development the male may bring more water stored in his cloaca if the puddle is evaporating; once the eggs hatch, the male will gingerly free the tadpoles from the sticky egg mass with his hind limbs; while the male had been tending to and guarding the egg site, the newly hatched tadpoles locate him and wiggle onto his back adhering to him by mucus secretions, where he then will carry up to 12 at a time to a more suitable water mass, such as accumulated pools in the center of bromelids; newly metamorphosed froglets are black with a pair of gold dorsolateral stripes down the back; in undergoing an ontogenetic color change, once the frogs reach maturity, the stripes disappear and the permanent adult color appears; considered one of the most intelligent of anurans; utilize strategy in hunting and rarely miss a direct strike with their long adhesive tongue; group members readily accept new juveniles from mothers of their group; captives recognize and pay more attention to their caregivers; during mating, emotion may even be observed with gentle cuddling and stroking between the mated pair; though there exists little predation, year round breeding and excellent parental care, this species is endangered due to habitat destruction and heavy collecting.

Endangered (3.1) IUCN

1. Maud Island frog ~ L. pakeka

Maud Island Frog
Family: Leiopelmatidae
Genus: Leiopelma
Species: Leiopelma pakeka

Length: 38.4-46.2mm/1.51-1.82in snout-vent length (SVL)
Weight: 6.00-10.75gm/0.21-0.38oz
Longevity: > 30 yrs; average life expectancy calculated at 33yrs
Name Origin: Leiopelma pakeka ~ Leios is derived from Greek meaning 'smooth'; pelma is also Greek meaning 'sole'; pakeka refers to Te Pakeka, the name given to Maud Island by the Ngati Kuia tribe; altogether means 'exclusive smooth skinned frog of Te Pakeka'.

Distribution: Endemic, but not confined to Maud Island of Marlborough Sounds in New Zealand; for species conservation purposes, individuals have been taken from the island and

2. Endemic range

translocated onto Motuara Island in 1997 where an established population is successfully growing; also, in 2006 21 individuals were placed in the Zealandia (formerly: Karori Wildlife Sanctuary) near Wellington, which is a predator proof small reserve; prefers cool misty surroundings within a temperature range of 8-14 °C/46.4-57.2 °F within the island's stable climate; is found in shaded areas of deep boulder banks on lower slopes under canopied coastal forest.

Description: This archaic little frog, along with the other 3 extant leiopelmatid species are living fossils, distinct from other frogs in having a direct ancient lineage going back 120 mya; 3 species just became extinct 1,000-2,000 years ago; primitive anuran traits still witnessed in L. pakeka are the retaining of tail muscles for wagging in tailless adults, an extra vertebrae making a total of nine and possess no tympanum (external eardrum); nostrils are nearer the eye than tip of snout; skin is relatively smooth with sparse scattered tubercles over the proximal portion of the thighs & shanks; the eye pupil is rounded; there are 4 toes on the forefeet with 3 on the hind feet; no webbing exist between toes; there are virtually no trait differences between sexes so are monomorphic rather than dimorphic as is the case in most frogs; this cryptic frog's coloration is well suited for camouflage in its native environment; dorsum is variable ranging from a uniform black and dark brown to light brown interspersed with irregular darker patching;

3. Note cryptic coloration

in the lighter skinned individuals, a black band extends from tip of snout to nostrils with a black line extending beyond the eye; ventrum side is variable in color ranging from contrasting light and dark blotching to a more uniformly pale or dark; may have the ability to lighten color during darkness as specimens observed at night appeared lighter than the same individual during daylight; appearing as morphologically the same (except for a couple of millimeters smaller), originally, L. pakeka was considered a variant of L. hamiltoni until late 1990s genetic studies deemed it as a separate species; later studies in the mid 2000s are now disputing this and putting into question as whether it should be listed as separate; there are no subspecies.

4. Lighter individual; note dark stripe

Diet: Diet consists of small invertebrates including insects, spiders and mites; there were no native predators on the island until the invasive Rattus rattus (black rat) & Mustela erminea (the stoat or short-tailed weasel) were introduced inadvertently by man.

Habits: L. pakeka is strictly terrestrial throughout all its life stages insinuating the frog line was first a land animal and later evolved in adapting to an aquatic lifestyle; relying on camouflage, when alarmed it will remain frozen in hopes of avoiding detection; it is a very clumsy hopper; when hopping it lands on its ventral side in 'belly-flop' fashion, recovers by pulling its sprawled legs back up into hopping position to repeat process; this is another hint that mastering of jumping came later in frog evolution; also possesses cartilaginous presacral ribs that later became vestigial in modern frog evolution; since there is no eardrum, a vocal sac never developed making the frog voiceless, although they can force exhaled air out into a high pitched squeak; L. pakeka has a tongue but cannot extend it out of the mouth; therefore, it must first lunge to

5. Froglets upon just hatching

capture prey with the mouth; communication is performed through chemical signaling rather than vocalization; the feces scent resulting from a male's high quality diet, attracts females; marked territory of no more than 5m/16.5ft radius is considered an individual's home range and it will stay put in this range for years; breeding occurs in December, in which is a warmer month for the southern hemisphere; 1-19 eggs are laid in a dampened or moist site on the ground as a gelatinous mass; the female leaves after ovipositing the egg mass under plant matter or rock cracks, but the male remains at the egg site guarding and tending to the eggs; there are no free tadpole stages as embryonic and larval (tadpole) development occur inside the egg; after a rather long incubation period of 14-21 weeks, the eggs hatch as miniature froglets with tails; the reason the tail remains is to allow more skin surface area to take up oxygen as the lungs aren't yet fully developed; once hatching occurs, the froglets crawl up the male's legs and onto his back where they will remain as inactive and not feeding until they are fully developed; once matured enough, the juvenile froglets leave their father's back and begin fending on their own; even though the eye pupils are round, typical of diurnal activity, this frog is nocturnal; though these frogs are fully terrestrial, they do require moist environments in combating desiccation and can swim; however, they swim awkward in alternating leg strokes that forces the head from one side to the other; this is not an energy sufficient method and by far the mastered swimming style of modern day frogs is far more thorough; though there are a few other frog species inhabiting New Zealand, they have been introduced; only the 4 extant members of the family, Leiopelmatidae are native to the islands that make-up New Zealand; due to its small native and unique habitat, this frog is listed as vulnerable.

Vulnerable (3.1) IUCN

1. Helmeted water toad ~ C. gayi

Helmeted Water Toad

Family: Calyptocephalellidae

Genus: Calyptocephalella
Species: Calyptocephalella gayi

Length: ♂ ~ 120mm/4.72in; ♀ ~ 12.60in snout-vent length (SVL)
Weight: Average: 500gm/17.64oz; females can reach up 3kg/6.61lb

Longevity: ~ 10-15 yrs

Name Origin: Calyptocephalella gayi ~ Calypto is an element used in many compound words of Greek origin meaning 'hidden' or 'covered'; cephalella comes from the Greek root word kephala meaning 'head'; gayi pertains to Claudio Gay Mouret (1800-1873), a French zoologist who was first to study Chilean flora & fauna; altogether refers to, 'Gay's shielded head frog'.

Distribution: Extent of range occurs between Coquimbo & Puerto Mott in southern Chile from 0-500m/0-1,650ft elevation; inhabits aquatic environments prefering lentic (standing or very slow waters) environments such as lakes and deep ponds; will note here that east of the Andes in southern South America, where climate is now cool and dry, a fossil of the extinct hyperossified, Calyptocephalella pichileufensis was discovered in strata of 47.46 mya when temperatures were much warmer during the Eocene.

Description: C. gayi, formerly known as Caudiverbera caudiverbera, was renamed by taxonomists C.W. Myers & R.B. Stothers in 2006, because they did not like the idea that Caudiverbera was based on a mythical species; why this was a problem, I don't know, for the rather large frog family, Hylidae is based on the mythical boy, Hylas; C. gayi was just recently removed from the family, Leptodactylidae and put under Batrachophyrnidae, but once again after reviewing prior karyotypic work, along with the rare genus, Telematobius, C. gayi was placed under the family, Calyptocephalellidae as the lone representative (monotypic) of the genus, Calyptocephalella; confused yet...; besides the size of this rather large frog, most

2. Close-up of head

distinguishing feature is the stark contrast with its smooth tight skinned head against a coarse bumpy loose skinned body making it appear as if it's helmeted, thus the common name; the frog is also colloquially called the 'wide mouthed toad,' for the mouth covers the entire perimeter of the head and can be opened wide to accommodate its ravenous appetite and facilitate its aggressive behavior; the snout itself is short and round; protruding eyes are tiny for its large size and exhibit bronze irises; tympanum is distinct on each side on back of head; the body is robust with dorsal skin possessing bumps that mass together forming ridges down the back in

3. Note dorsal ridges down back

a lot of individuals; relatively short forefeet are unwebbed while large hind feet are close to halfway webbed; overall coloration is a dull brown to greenish with fainter irregular markings; possesses a grayish white ventrum; there are no recognized subspecies.

Diet: Consumption consists of other frogs, but will take anything that moves near it, such as aquatic life (insects & their larvae, fish), smaller mammals and birds; natural predators of this frog as an adult are insignificant except for human's harvesting and consumption of the frog, which does take its toll on certain C. gayi populations; fish will take earlier stage tadpoles.

Habits: This frog is one bad boy; when cornered it is aggressive in lifting up the body as it inflates its lungs to maximum with air and in opening its mouth will lunge to bite its intended intruder; this is followed by loud sequences of modulated cries with mouth agape in contrast to harmonious mating calls with mouth closed resonating off one repeated call; with short but

4. Note folding of legs

jagged cusped teeth on upper jaw, it can inflict a painful bite; aggresive behavior and cry frequencies occur with both sexes; the modalities of aggressive behavior in these frogs are considered a resultant of a long independent evolution; with a vertical pupil, this frog is active both night & day; prefers to lie in wait and ambush any animal that comes into range; as most frogs, detects prey from movement; found in the foothills of the Andes, this frog prefers cooler water temperatures where it remains most of the time uncommonly going ashore; has a habit of folding its hind legs inward over the anterior portion of the dorsum when at rest or in hiding; breeding season occurs from September to October where male emit their mating calls of 'oouu'; at this time males also produce a nuptial protuberant pad on the fore toe that would represent a thumb; the nuptial pad is more gaining a firm grip when clasping the female during amplexus; the female lays her clutches in the shallows with muddy bottoms; fertilized eggs are oviposited in clumps consisting of 800-

5. Tadpoles (larvae) held for size ratio

1,600 in a gelatinous mass; hatching occurs in ~3 weeks where the tadpoles migrate and aggregate to the muddy bottoms in around 1m/3.3ft of water with plenty of aquatic vegetation; the larvae grow very large up to 150mm/5.91in in length and weigh 30gm/1.06oz; the giant tadpoles appear as Gymnotiformes (knife fish); it takes a full two years for the tadpoles to fully metamorphose into froglets and throughout the aquatic metamorphic stage remain rather slow and poor swimmers; the newly metamorphosed froglets measure 22mm/0.87in; primarily due to drainage of habitat, introduced pesticides & pollution; introduction of certain fish as trout consuming larvae and the trapping & hunting by humans for consumption populations are in decline; where it was once abundant in natural ponds and small lakes near towns, this frog is practically absent now; 30% of its population has decreased within the last decade; due to continual population declines, it is now listed as threatened.

Vulnerable (3.1) IUCN

1. White-lipped frog ~ L. fragilis

Mexican White-lipped Frog

Family: Leptodactylidae

Genus: Leptodactylus

Species: Leptodactylus fragilis

Length: 3.8-5.1cm/1.5-2.0in

Weight: inferred: 5.96-11.81gm/0.25-0.43oz

Longevity: Unknown

Name Origin: Leptodactylus fragilis ~ Lepto is Greek meaning 'slender' and daktylos is also Greek meaning 'digits'; fragilis is Latin for 'fragile'; altogether means, 'frog with thin and fragile toes'.

Distribution: Range occurs marginally from the northern end in 3 extreme Texas counties of the lower Rio Grande Valley; found in one locality of Cameron & Starr counties and in 2 localities of Hidalgo county; range continues fully down throughout Mexico's Caribbean coastline and up to its Pacific coastline from the extreme southwestern end up to Veracruz; is found in all 7 Central American Countries; in extreme southern range, becomes marginal again with intermittent populations found in Colombia & Venezuela; prefers semi-dry conditions in semi-tropical and tropical forests, second growths and low lying grassland with structures like ditches and ravines that semi-permanently or seasonally collect rainwater.

Description: There was some confusion on the naming of L. fragilis, so in 1978 the name was changed to the same species as, Leptodactylus labialis in reference to the white striped lip; however, in 2002 it was verified that Leptodactylus fragilis was indeed the proper name and

2. Distinguishing white lipped stripe

was switched back as the valid species name; not sexually dimorphic, except male head may be wider; most distinguishing feature on this small frog is the white or beige lipped striping and long spindly toes on each foot; has a defined longitudinal line along rear surface of each thigh; skin of tarsus and soles of feet possesses prominent white tubercles; has a pointed snout on a rather large head; vomerine teeth are present in transverse series located entirely behind the choanae (paired openings between nasal cavity and nasopharynx; with hind leg osteology, are efficient leapers; distinct typanum present; dorsolateral fold exists along torso; circular ventral disk is present on ventrum; coloration varies between individuals with most being gray with the rest showing a reddish bronze or muddy brownish green dorsum; dark blotching appearent, but not confluent; currently there are no subspecies.

3. Brownish-green dorsum

Diet: Is a generalist/oppotunistic feeder, almost exclusively consuming small arthropods consisting of mites, ticks, spiders, diplodans (millipedes), insect larvae and insects, in particular coleopterans (beetles) and blatiids (cockroaches); a very small percent of diet includes molluscs; predation on this frog abounds from other larger frogs eating L. fragilis to reptiles, birds, bats and the jaguarundi; large water beetles will take tadpoles.

Habits: These frogs are primarily terrestrial and are nocturnal lying in self-dug burrows just underground during the day; they forage nocturnally in open areas; where ranges overlap, Leptodactylus columbiensis is more common in wetter habitat, where L. fragilis is higher in

4. A Belizean reddish bronze specimen

number in the drier zones; something we will call here, 'heterospecific eavesdropping' is evident with the predator alert male calls of L. fragilis; other male frog species will ignore other male frog species' predator calls, but pay attention and respond to L. fragilis' alarm calls; Where Physalaemus pustulosus & Physalaemus enesefae are closely related and have similar calls, they both will respond rapidly to a L. fragilis male's predator alarm call more so than they would to their own respective species call; this attendance to predation related calling consisting of sympatric, but heterospecific communities appears to be commonplace among these 3 frog species survival strategies; male L. fragilis calls are extremely dissimilar to the other species' frog alarm calls, but the interspecific eavesdropping must be successful in natural selection between varying frog species in predation cues, in particular when it relates to frog eating bats; predatory bats seem to hone in on P. pustulosus' chucking predator alarm cues, so that is most probably why P. pustulosus and P. enesefae (with similar chucking predator cues), relies more on L. fragilis to

5. Male excavating nesting site

be the watch dog of the frog community; sexual maturity is reached when males achieve a length of 36mm/1.42in and for females, 40mm/1.58in; breeding takes place during heavy rains; the male excavates a brooding chamber in moist ground and begins his mating calls of frequency ranges in the northern portion in Texas at 600-1200 hertz (hz) and of 1000-2200 hz in its southern range; frequency is modulated with intermittant pulsating and nonpulsating portions; brooding chambers are usually located under clumps of grass, dirt clods, logs or other natural ground litter; when female is attracted mating occurs externally and the eggs are laid in the nest chamber encased in bodily secretions whipped up into foam to avoid dessication; clutch size varies with up to 86 yellow eggs laid in northern range in Texas and between 25-250 eggs in more southern ranges; egg development takes place in the foam nest where tadpoles hatch as larvae into the chamber; from rains, larvae are swept away to runoff areas such as low lying grassland or bodies of water such as oxbow lakes and ditches; northern larval stages take 30-35 days to fully develop, where southern ranges, full metamorphism is achieved in less than 2 weeks; currently, L. fragilis is not threatened; except for possible extirpation from Texas localities due to heavy use of organophosphate fertilizers, populations appear stable.

1. Edible frog ~ P. esculentus

Edible Frog

Family: Ranidae

Genus: Pelophylax

Species: Pelophylax kl. esculentus

Length: ♂ ~ 5-9cm/1.97-3.54in; ♀ ~ 6-12cm/2.36-4.72

Weight: 5-12gm/0.17oz-0.4oz

Longevity: 5-15yrs

Name Origin: Pelophylax kl. esculentus ~ Pelo is Latin for 'around the' in referring to muddy shore; phylax is from the Greek root word, phylakos meaining 'to guard'; esculentos is Latin meaning 'edible'; altogether refers to 'edible guardian frog of muddy bank'; kl. is the abbreviation of the Latin word, klepton which means 'hidden' or 'concealed'; the reason it is included in the scientific nomenclature is to designate that this species is not pure; in being a diploid hybrid, but fertile species, it retains 'concealed' genetic material within its DNA of another species' chromosomes; P. esculentus must have an exchange of other species' genes in order to survive as fit.

Distribution: Found across all of central Europe from its northern range in Germany & Estonia to the southern range of southern France, northern Italy and Croatia; natural migration population pockets exist in Sweden & Bulgaria and introduced populations in England & Spain; prefers remaining near shorelines in sunny positions of ponds, brooks, canals and marshes with vegetative growth.

Description: P. esculentus (formerly classified as Rana kl. esculenta) is a fertile hybrid of the pool frog (Pelophylax lessonae) and the marsh frog (Pelophylax ribibundus) and by geological standards, is a recent species to arrive on the scene; during the last glacial period, ~20,000 years ago the population of one common ancestor for P. lessonae & P. ribibundus was split apart due to glaciation diverging the ancestor's population enough that morphologically and physiologically the two population halves became distinct species as the pool frog and marsh frog; once glaciers receded ~10,000 years ago, the 2 species were still genetically similar enough to interbreed successfully with P. esculentus as the resultant hybrid offspring; as a medium sized frog displays sexual dimorphism with female as slightly larger; males possess vocal sacs on sides of each cheek positioned just behind the mouth angle, while females do not; only males have extra skin patches (nuptial pads) on their forefeet; males have developed metatarsal spurs for clasping during amplexus while limbs are slimmer in females; irises of eyes

2. Male displaying vocal sacs

are golden yellow in both sexes; skin is permeable; cohabiting the same environments as the parent species, the edible frog is commonly mistaken as the pool and marsh frogs; though intermediate characteristics of parent species are retained by the edible frog, differences are distinct; marsh frogs are more robust, warty, have hind legs that extend well beyond snout of the head, dark gray vocal sacs and has the harshest mating call that is more rapidly delivered; the pool frog is the smallest and most compact of the three frogs, hind legs do not extend to or past the snout and has white vocal sacs; the edible frog has hind legs that extend just to the snout and has light gray vocal sacs; overall coloration of P. esculentus is from greenish, olive green to grayish green; dorsum has light patches of dark browns usually with a light yellowish green mid-dorsal line going down the spine of the back; white ventrum speckled with dark spottings; males become much more greener during mating season; there are no designated subspecies.

Diet: Is an opportunistic predator, preferring to for prey to come by while lying in wait concealed along the shoreline; consumes all invertebrates including molluscs, insects & worms and will also attempt to eat small birds & mammals; it is known as well to be cannibalistic, consuming smaller frogs of other species and it own species; this frog will also leap to capture flying insects such as dipterans (flies), hymenopterans (wasps/bees), orthopterans (grasshoppers/crickets) & odonates (dragonflies/damselflies); will venture out on land around 500m/1650ft from shore or hunt in water for prey if none aren't available along the shoreline; tadpoles are essentially herbivorous eating aquatic vegetative matter, but will consume micro-

3. Cannibalism

organisms; Primary predators are shorebirds, owls and grass & water snakes; P. esculentus supplies the frog legs that are considered a culinary delicacy in France and other countries; where present, fish will take tadpoles as a meal.

Habits: Unlike most, this frog is diurnal preferring to hide motionless near or in the water during darkness; in its more northerly clime ranges, hibernates from October to March; in hibernating, takes to land burrows, but hibernating in the water bottoms have also been observed when coexisting with P. ribibundi; when alarmed will instantly jump into water; if caught will emit loud screeching sounds; mostly seeks the southern banks receiving the most sunlight and vegetative growth; are territorial, so therefore are mainly found individually and guard their turf by emitting a rising cadence of 'qwarks' towards an intruder; males though, do congregate

4. Pelophylax kl. esculentus

in still waters when they begin competing for females during mating season and sometimes in high densities; males become aggressive toward one another and may eject liquid jets from their cloacas; breeding season occurs primarily during May, but can extend into June; mating coincides with the breeding period of P. lessonae & P. ribibundus; when one male begins the mating call, all others begin to chorus in; females only become aroused when she hears parental species males chorusing in with P. esculentus males; when females are attracted, she enters the water; once a male recognizes a female by sight, he quits singing and swims over to her creating an aggregation of males surrounding the female; the group of males then lead the female to an area of dense vegetation where all the males begin their courtship singing; the female prefers the loudest and as judge picks her favorite; the winning male then mounts her and they begin to perform amplexus; in this process, she will oviposit a small clustered ball of eggs that he fertilizes as they exit her cloaca; the female interrupts her egg laying until the male is finished fertilizing, then they both repeat the process until she has laid between 3,000-10,000

5. P. esculentus tadpole

eggs; coloration of eggs are brown on top and creamy to white on underside; eggs take 2-3 weeks of incubation and hatch in June as tiny 5mm/0.20in tadpoles; although in last stage of larval metamorphosis, the 7cm/2.76in tadpole turns into a 20mm/0.79in young froglet; to be noted here, where cohabiting with parental species, the female P. esculentus prefers the male P. lessonae when present and the male P. ribibundus when present over the male P. esculentus; deformities can result such as an abnormal sacral region when both parents are P. esculentus; this complex hybridogenetic form of reproduction is very rare in nature and together with the 2 parental species and P. esculentus diploids, hybrid populations may also contain triploids with a cell containing 3 sets of chromosomal information material; though by law, there are no protections in the countries where the edible frog was introduced, the population has slightly increased outwards from the introduced ranges; though slightly uncommon in its south easterly range, populations as a whole appear stable and are not threatened.

1. Bullfrog ~ R. catesbeiana

American Bullfrog

Family: Ranidae

Genus: Rana

Species: Rana catebeiana

Length: Females typically larger, but both sexes can reach 15.2-20.3cm/6-8in SVL

Weight: Up to 750g/1.7lbs

Longevity: 8-10 yrs in wilds; Record in captivity: 16 yrs

Name Origin: Rana catesbeiana ~ Rana is Latin and probably mimicked the sound of how Romans heard and interpreted the frog's call; catesbeiana honors Mark Catesby (1682-1749), who was one of the first naturalists to study the flora & fauna of Virginia and produced, 'Natural History of Carolina, Florida and the Bahama Islands'; altogether refers to, 'Catesby's croaking frog'.

Distribution: This North American frog is native to central & eastern U.S. extending into the southern portion of Canada in Quebec & Ontario; has been introduced into western portions of U.S. creating patchy populations from state of Washington down through Oregon, California, Arizona, New Mexico, extending into northern Mexico, back up into Colorado, Wyoming, Utah, Nevada & Idaho; virtually all of Texas is inhabited once introduced into the extreme west; prefers the banks of still waters such as ponds, swamps and lakes with plenty of vegetation.

Description: R. catesbeiana is the largest frog native to the continent of N. America; bullfrog skull is highly fenestrated (perforated); continuous rows of tiny spiked teeth exists on the maxilla & premaxilla with one pair of vomerine teeth on palate; mandibles support no teeth; though skeletal bullfrog structure represents a true tetrapod, as in all ranids, it bears no ribs;

2. Osteology of R. catesbeiana

each vertebra has extended pedicles; except for longest fourth toe, hind feet are fully webbed; dorsolateral folds end near tympanum by folding around top and body side of the eardrum; this species shows sexual dimorphism; besides females being larger, there is a prominent external tympanum that is the same size as the eye in females and a bit larger in males; throats of males are yellow where females are a creaminess white; the eye's iris is either golden or reddish bronze; overall coloration is highly variable from a lime green to a gray net-like pattern over a green ground color and to a greenish brown; back & sides may possess irregular black spotting; the forelegs & hind legs are barred or spotted with black; ventral underpart is normally an off-white mottled with gray or yellow; there are no designated subspecies.

3. Note webbing of hind foot

Diet: Is an opportunistic ambush feeder and will consume any animal that moves near it that can be overpowered and swallowed; included in the frogs diet is an array of animals from insects, worms, snails, crayfish, small reptiles such as turtles & snakes, small birds and small mammals; even other frogs and bullfrog juveniles are taken; the appetite is voracious and is constantly in wait of some animal to eat; where most other large frogs can't, the bullfrog is adept and quite capable of capturing prey underwater; the tadpoles are primarily herbivorous; just as numerous is this frog's prey, so too are its predators; larger turtles, snakes, fish, mammals and birds greedily consume this frog; it is eaten by man throughout its range and in fact the western introductions were initiated solely for the purpose to set-up populations for human consumption.

4. Note male features

Habits: Is crepuscular, but mainly nocturnal; is solitary and territorial, rarely encountering one another except during breeding season and males fighting over territory; preferring to remain along the water's edge concealed in vegetation lying in wait for any potential prey, it expends little energy; if a threat appears, it first will choose to flatten body to ground; if the threat ensues in becoming too close for comfort, it will either leap into the water or into the bank's surrounding vegetation; the frog is able to leap a good distance of 1-2m/3.3-9.9ft with its long powerful hind limbs measuring 17.8-25.4cm/7-10in; also uses hind limbs' webbed feet to proficiently and rapidly swim; in kicking out, uses hind limbs only in swimming holding forelimbs out of the way against the sides of body; hibernates throughout its range during the colder months by burrowing into the mud beneath the water;

5. Swimming style

R. catesbeiana breathes via 3 methods: subcutaneous through the skin, gas exchange through the moist mouth and with a pair of lungs; being highly aquatic it is rare to see this frog venture any distance from its water source; when capturing larger prey such as a vertebrate, it will use its forefeet to help push the prey further down into the mouth; if the prey is struggling or fighting back, the bullfrog will jump underwater in attempts to subdue and drown the prey; in knowing that its vertebrate prey can breathe only in atmospheric air, it would be interesting to know if the frog does this instinctively, or it is a learned behavior; breeding may take place anytime from February to October, but is kicked in high gear from late May up into July; males first collect vying for the best egg depostion spots; males possess a single internal vocal sac that resonates bass tones; to attract females the male begins emitting his famous deep 'jug-o-rum' song that can be heard 0.40km/0.25mi away; most say the 'jug-o-rum' calling is reminiscent of a bull roaring, hence the colloquial name; once a female is attracted, the male initiates

6. R. catesbeiana variable tadpoles

amplexus by mounting her and grasping her with his forelimbs posterior to hers; the female will oviposit 12,000 eggs while sometimes up to 20,000 over the nesting site the male picked and defended from other male rivals; as the eggs exit the female's cloaca, the male releases his sperm externally fertilizing the eggs; the eggs are laid in a water insoluble film causing the whole egg mass to float on the surface; incubation lasts within one week and dependent on temperature, hatch in 4-5 days; most tadpoles after hibernating, fully metamorphose into froglets in their second year, but some remain as tadpoles until their third year; tadpoles can be large reaching lengths of 125-150mm/4.92-5.91in; the larger the tadpole, the larger it will be as a frog; tadpole coloration is as varied as adults and appears to be dependent on geographical range; folks that consume the frog legs and upper rear back purport the bulky muscles taste a little like chicken; 'frog giggin' as the term of hunting bullfrogs is called, is conducted at night with a spotlight and trident; guided by the sound of the rumping bullfrog, the light is shone on the eyes temporarily immobilizing the frog, then after slowly moving within reach, the bullfrog is gigged with the forked trident; regardless of the pressures of human consumption, R. catesbeiana is not considered as threatened.

1. Owl Frog ~ H. australiacus

Owl Frog

Family: Myobatrachidae

Genus: Heleioporus

Species: Heleioporus australiacus

Length: 60-100mm/2.36-3.94in

Weight: > 100gm/3.53oz

Longevity: Unknown, but possibly at least up to 10 yrs due to size and slow metabolism

Name Origin: Heleioporus australiacus ~ Heleio is Greek for 'marsh' while porus is also Greek meaning 'opening' in reference to pores; australiacus refers to 'pertaining to Australia; altogether refers to, 'porous marsh frog of Australia'.

Distribution: This endemic species is confined to two sub populations with range constricted to the eastern slopes of the Hawkesbury Sandstone region in southeastern Australia's coastal range; is found from near sea level to 1,000m/3,300ft in elevation and as far inland as 100km/62.14mi; northern population inhabits the dry Sydney basin while the more southerly population inhabits wetter montane woodlands in Jervis Bay; there is a 100km/62.14mi distance between the 2 disjunct populations; there may be specimens in between that have not been detected due to little & limited studies; though species lives in two different habitats, prefers areas dominated by sandstone plains in bushlands or dry & wet sclerophyllous woodlands.

Description: Also known as the 'giant burrowing frog,' this species is one of Australia's largest frogs; is rotund with portly but stout limbs; there are traces of webbing on the hind feet with no webbing on the forefeet; metatarsal tubercle is unpigmented flesh; dorsal side is warty with ventral underside being slightly smooth to granular; parotoid and tibial glands are absent; snout is very short and rounded; tympanum is distinct and ~ same size as eye; Heleioporus

2. Note short snout

australiacus exhibits dimorphism with the male having the sides, arms and portions of the back covered in black warts capped with fleshy but hard spines; pupils are vertically elliptical with silver irises; in both populations, there is great variability in coloration; dorsum ranges in color from light brown to a rich deep chocolate to black; sides range from a light gray to bluish usually with random yellow warts; most of the time there is a yellowish splash underneath the front limbs; behind the mouth on each side a row of lumpy yellow spots are normally present; ventrum is an off-white usually with some brown on the throat area; tadpoles are from gray to brown to ash black; tadpole physique is much different than typical tadpole features in being flattened with a disc shaped head that might remind one of limulids (horseshoe crabs); though the species lives in two different habitats it has been determined that it is indeed one species conforming and acclimating to varying environments, instead of those environments catering to two species specialized needs; although a small gene study did lend towards two distinct species, more work needs to be done to verify; therefore, currently there are no subspecies.

3. Note: silver iris & tympanum

Diet: Is considered a generalist predator consuming invertebrates, including terrestrial & flying insects, centipedes, spiders and scorpions; ants appear to make-up the bulk of its diet; foxes, laughing kookaburras (Dacelo novaeguineae) and the red-bellied black snake (Pseudechis porphyriacus) are natural predators of the frog primarily during its breeding season when the frog is more out in the open; far more mortality rates are due to domestic cats and road kill.

Habits: The occurrence of this frog is limited to an 80,000 squared km or 49710 miles squared; even within that range they are located usually when breeding in the open; in being a burrowing frog, are rarely encountered; when located and cornered will go into a defensive display in stiffening its legs raising the body while also inflating with air to appear larger; in

4. Defensive posturing

5. another defensive pose

this defensive pose, a creamish and toxic irritant is exuded from skin secretions all the while the frog is emitting a mournful cry; if that doesn't alleviate the threat and an encounter pursues, males will powerfully thrust their forelegs utilizing the spikes as a piercing and rasping weapon on whatever gets between its forelegs, whether it be a male opponent, a potenrial predator or a misguided human finger; is nocturnal spending most of the daytime avoiding detection burrowed underground; also burrows during unfavorable conditions and to hibernate & aestivate; heavy rains will bring them above ground; will range widely moving up to 300m/990ft from its home range; usually burrows its own hole, but taking up other animal burrows is not uncommon; excavates with rear legs reversing in a revolving manner until whole body is covered there are 3 types of burrows: 1) temporary burrows for hiding away which are shallow just beneath the surface; these are used for avoidance of predators & fires; 2) aestivation & hibernation burrows are much deeper and usually located in stream banks or near ponds; other animals may share this burrow; 3) breed

6. H. australiacus tadpole

burrows have two openings located in banks of slow moving creek lines & pond areas; male calls from within or near these breed burrows or within vegetative debris initiates female response; call is much like an owl's hooting o,f "ou-ou-ou-ou-ou" lasting up to 2 second intervals; thus the common name of owl frog; males call year round, but are most active in the months of December through February; once the female is attracted, amplexus occurs within the burrow chamber with the male utilizing his spines to clasp the female; eggs are simultaneously deposited and fertilized in a moist foamy mass inside the chamber; foamy egg clutches contain between 40-1200 eggs; hatching of the eggs and early development begins within the breed burrows, or vegetative debris; coinciding with rain, later, a subsequent heavy rain will flush out the tadpoles into the more permanent body of water they were first laid near; tadpoles take from 12 weeks up to 6 months for full metamorphosis with the later developing tadpoles hibernating first before fully developing in warmer weather; due to development of their confined habitat, H. australiacus is listed as threatened.

Vulnerable (3.1) IUCN

1. Common midwife ~ A. obstetricans

Common Midwife Toad

Family: Discoglossidae

Genus: Alytes

Species: Alytes obstetricans

Length: Up to 55mm/2.17 SVL

Weight: 8.40-9.65gm/0.30-0.34oz

Longevity: Ave: 5yrs

Name Origin: Alytes obstetricans ~ Alytes is Greek meaning 'entwined' or 'linked'; obstetricans is Latin for 'midwife' for the reason that during mating, the male inserts a toe into the female's cloaca to aid her in egg laying; altogether refers to the 'linked midwife toad'.

Distribution: As native, this frog occurs wholly or in parts of 9 European countries in Belgium, France, Germany, Italy, Luxembourg, The Netherlands, Portugal, Spain & Switzerland; has been introduced into the United Kingdom; breeding & larval water habitat preference varies greatly from ponds to creeks to slow rivers ranging from sea level to an elevation of 2400m/7,920ft in the Pyrenees, Massif Central & the Alps, but is uncommon above 1500m/4950ft; prime adult land habitat is stony or sandy slopes and embankments; prefers ground with lots of sun exposure and little vegetation; has established large colonies in gravel or clay pits.

Description: This species is a small, but stocky frog; sexual dimorphism is evident with female normally a bit larger than males in a population, though both sexes can reach 55mm SVL; there are other minor differences between sexes such as distance between nostrils than eyes are large with vertically elliptical pupils; parotoid glands are present but small, tympanum is also small but usually visibly apparent; dorsal skin is warty on body & legs; there exist glands complexes located in underarms & ankles; often reddish warts extend in a line from the tympanum to loin area; overall body coloration varies from a dorsum dark gray to yellowish orange with some specimens being light green with darker green warts; Ventrum is a dirty white with throat & chest area often spotted with gray; There are currently 4 recognized subspecies:

2. Female, note row of reddish warts

A. o. obstetricans, A. o. almogavarri, A. o. boscai & A. o. pertinax.

Diet: This little frog feeds on smaller invertebrates with half of its diet consisting of spiders and beetles; food intake is rounded off with other terrestrial insects, flying insects, pillbugs, slugs, snails and worms; when breeding the frog does not eat; tadpoles are primarily herbivores eating plant and organic material; once metamorphosing into a froglet the intestines shorten aiding in converting the animal into a carnivore; primary predators that prey on adults are fish in particular introduced fish) during males release of hatching eggs into water; other than fish natural predation is low due to toxins secreted by parotoid glands and dorsal warts; certain individuals of bird species will take the frog though; tadpoles are consumed by water bugs, fish & the viperine water snake, Natrix maura.

3. Male with eggs inflating

Habits: As a lot of frogs in defense, this terrestrial species also inflates the front portion of its body with air, accommodated by the tucking in of their limbs underneath body; inflated posture is held up to 5 seconds; lives on land as an adult, but usually no more than 100m/330ft from water permanence; in hottest months to retain moisture, though sometimes occupying other animal dugouts will usually dig a burrow themselves with their paunchy forelimbs and hideout there during daytime heat; the high pitched whistle-like calls are described as, 'poo-poo-poo' and are reproduced every 1 to 3 seconds; males don't sound until after March into late summer, though in colder regions, may sing until November; this frog is best known for its male paternal care of eggs; reproduction takes place in spring and

4. Male with eggs

summer months; females seek out the male inviting him to mate by stamping her forefeet and gently nudging him; male will then perform amplexus using his hind toes to stimulate her cloaca; within 10-20 minutes he will squeeze female's flanks causing her to stretch her legs and begin laying eggs in a clutch of around 45-50; once inseminating the eggs, the male will then distends the egg mass with his hind legs, ply them alternatively to his body and extend the string of eggs until they wound around his ankles; he will then utilize his front legs to ensure proper adherence of the eggs to his body. females can produce up to 4 clutches of eggs per breeding season and male will mate with other females to add another egg clutch, males can carry up to 3 clutches and with eggs from previous mating twined around his legs and back portion will carry up to 150-170 eggs; females prefer mating with a male is not already carrying an egg clutch and if she does mate with an egg bound male may release less eggs than what

5. Egg bound male in day burrow

she is actually carrying; males will carry the eggs for 3-6weeks until they hatch, keeps them from ground contact by keeping hind limbs slightly elevated off ground; during heat of day to avoid egg desiccation, remains in burrows until coolness returns; if the eggs are in danger of drying, will take a dunk in the water to refurbish moisture; depending on climate regions; some males are seen carrying eggs at the end of August; the toxin gland secretions from his warts and parotoids help protect eggs from infection; once the male identifies particular movement from within the egg, it signals the time for release of eggs; the male may pick any permanent or suitable temporary body of cool water that is

6. A. obstetricans tadpole

stagnant or slow moving where he releases the eggs into the water; within minutes afterward the eggs hatch as tiny larvae 10-17mm/0.39-0.67in long; tadpoles lite speckled brown color creates good camouflage against muddy and sandy water bed; autumn tadpoles will hibernate before turning into froglets the next spring. hibernating tadpoles become large averaging sizes almost as large as the adult frog at 50 mm/1.97in; in exceptional cases there have been hibernating tadpoles observed reaching 100mm/3.94 in length; once tadpoles fully metamorphose, they turn into froglets measuring 12-15mm/0.47-0.59in, then leave the water; though introduced fishes have decimated populations along its northern border range and there have been mass die-offs due to chytrid infection, the species in general is not listed as threatened.

1. Ranita de cristal ~ C. ilex

Ranita de Cristal (Crystal Frog)

Family: Centrolenidae

Genus: Centrolene

Species: Centrolene ilex

Length: 28-37mm/1.10-1.46in SVL

Weight: Publishing resources unavailable; ~3.29-8.67g/0.12-0.31oz

Longevity: Unknown

Name Origin: Centrolene ilex ~ Centro & lene are both Latin meaning 'center' & 'easy' respectively and is in reference to ease of use and positioning of humerus spine in adult males; the specific name, ilex is Latin for 'holly plant,' but in this case is to honor herpetologist, Priscilla Hollister 'Holly' Starrett; altogether refers to 'Holly's centrally located for ease of use humerus spine frog'.

Distribution: Occurs in isolated pockets of known populations in Colombia, Ecuador and the Caribbean slope of Panama, Nicaragua & Costa Rica; originally was a species of South American landmass that spread out into Central America once Isthmus of Panama land bridge was exposed during Pliocene 3mya; ranges from almost sea level to montane subtropical jungles in elevations up to 1420m/4686ft; is evident in plant life along side streams and small rivers.

Description: Was formerly listed under, Centrolenella but genus was broken up into other genera; some taxonomist publishers list this species under the genus name, Sachatamia; as far as glass or ghost frogs go, this species is medium-large sized and is largest glass frog in Central America; most distinguishing feature is the forward pointed eyes sitting on top of the

2. Note distinguishing eyes

head; iris is silver with reticulated black markings; pupils are horizontally elliptical; prominent nostrils are are protuberant resting on a bony elevated ridge; exterior tympanum is evident but indistinct; vomerine teeth are present in transverse rows the choanae (openings between nasal cavities & nasopharynx) and are medially separated; limbs are spindly with extensive webbing on hind toes, while webbing is only evident on outer toes of forelimbs; toe discs are truncate; sexual dimorphism is evident with females reaching 37mm/1.46in when males only reach 28mm/1.10in; only adult males have a humeral spine embedded within the foreleg musculature; males also possess a white nuptial fore thumb pad located on dorsal and lateral surface; located on dorsal and lateral surface;

3. The more rare spotted specimen

dorsum and limbs are a leaf green with a whitish rim trimming the mouth; feet are yellowish green; ventral is transparent with no pigmentation revealing internal organs and eggs encased in females; digestive tract is also unpigmented, but a white peritoneal sheath covers internal organs; the bones appear dark green; white pigmentation may be evident on flanks; on rare occasion, white dots are evident on dorsum; there are no current subspecies listed.

Diet: Adults consume small terrestrial or aerial invertebrates such as insects and spiders; tadpoles graze on alga and decomposing organic debris; predation on this frog ranges from snakes, birds, arboreal mammals, larger tree frogs and fish consume tadpoles; below 500m/1,650ft frog-eating bats can be a predator distinguishing the frog's call from poisonous tree frogs.

Habits: C. ilex is nocturnal and arboreal spending the daytime hunkered down sleeping on leaves near streams; has the ability to change its dorsum green color to match exactly the green

4. Note white sheath

shade of leaf its resting on; toe bones are T-shaped to support the surface area of wide toepads used to adhere to vertical substrate; with a distinct neck and rounded head along with directly forward positioned eyes, this frog most likely has stereoscopic vision for maneuvering in lower canopy; males are territorial and will wrestle with another male intruder once entering his designated leaves' zone; it was originally theorized the humeral spines were utilized to clasp the female during amplexus; but further observations concluded that the spines are used in male combat where the wrestling opponent is pinned between the spine and foreleg; during the breeding season, males, facing towards tip

5. Translucent ventrum showing eggs

of leaf it is on, calls in repetitive single high pitched 'clicks;' click intervals are repeated every several minutes; once a receptive female arrives, amplexus takes place; eggs are externally fertilized; the female deposits her darkened eggs on the top surface of an overhanging leaf above running water; the egg clutch is encased in a clear & wet gelatinous mass; in laying eggs on plants, the clutch is less vulnerable to predation than those laid directly in water; to keep parasitic fly larvae from attacking egg clutch, parents may guard the eggs consuming or chasing away any flies that might lay their eggs on the egg clutch; upon hatching, the tadpoles fall off into the water and immediately going into hiding within the stream bottom's pebbles; the tadpoles are streamlined and elongated with efficacious tails and able low fins geared for faster flowing waters; though in late 1980s this species' Costa Rican populations were decimated after being infected with a primitive water borne pathogen (chytrid fungus) it is not listed as threatened; but, continual deforestation which is heaviest along stream lines where this frog lives may indeed threaten the species; if overhanging plants along streams are greatly reduced, C. ilex will become extinct as its egg laying strategy will be taken away.

1. Couch's Spadefoot ~ S. couchii

Couch's Spadefoot Toad

Family: Scaphiopodidae

Genus: Scaphiopus

Species: Scaphiopus couchii

Length: 5.6-8.8cm/2.25-3.5in SVL; ave: 6cm/2.36in

Weight: 17.18-33.13gm/0.61-1.69oz

Longevity: Up to 13yrs in the wild

Name Origin: Scaphiopus couchii ~ Skaphis is Greek for 'shovel/spade' while pous is also Greek for 'foot';couchii honors the zoologist, Darius Nash Couch (1822-1897) who first discovered this frog; altogether is in reference of, 'Couch's spade footed frog'.

Distribution: Is a N. American frog; range extends from all of central-west Texas into southwestern Oklahoma for northernmost boundary and down into the northern border area of Mexico westwards through southern New Mexico, southern Arizona ending in southeast

2. Prime breeding habitat

corridor of California & northeast Baja; found in a variety of arid habitat suitable for burrowing including desert, scrubland, shrubland, mesquite savannas; deciduous woodlands and in rocky outcrops where sandy soil is available.

Description: Formerly listed under Pelobatidae; due to recent mtDNA analysis, phylogenetic relationships proved Eurasian spadefoots are a sister group to megophryrids and are not monophyletic with N. American spadefoots; Eurasian spadefoots remained under Pelobatidae, while N. American spadefoots were given the new family listing, Scaphiopodidae over the two genera listings of Scaphiopus & Spea; spadefoots are only commonly named toads because they appear physically as toad-like, but they are truly not toads; physical similarities simply

3. Note spade located on hind feet

are convergent evolution as there are actually many morphological, anatomical and physiological differences between the 2 groups; Most striking feature of this frog are the black keratinized sickle-shaped spades located on each hind foot; unlike toads, spadefoots lack a raised cranial boss across eyes; possess small dorsum warts, but skin is smoother and wetter than true toads; there is an absence of parotoid glands in spadefoots; osteological differences is the coccyx of spadefoots is fused to the sacral vertebra with the sacrum having expanded diaophyses (a vertebra transverse process that is an outgrowth of the neural arch on the dorsal side); S. couchii has 8 presacral vertebrae that are procoelous (where vertebra is concave anteriorly & convex posteriorly); eyes are vertically elliptical; hind feet are nearly fully webbed where is mostly absent on forefeet; Sexual dimorphism is evident with females being larger and more marked; it is common for male individuals to be typically patternless; coloration is variable from green to greenish yellow to brown yellow; dorsum is mottled with dark green, brown or black markings being more distinct in females; ventrum is a pale white in both sexes; tadpoles are iridescent with a golden sheen over a coppery bronze hue; there are no subspecies currently listed.

In above photos, note variable coloration in individuals & varying color in male and female during amplexus.

Diet: This frog, when on surface in spring and summer will virtually eat anything that moves and can be stuffed down its throat; primary food are invertebrates such as insects and arachnids, snails, but in particular winged termites that begin to swarm when the frog comes out of hibernation; termites are an essential diet which are high in protein and fat content; good for a frog when inactive in hibernating most of year; tadpoles are omnivores consuming a variety of water borne organic matter from humus rich mud, algae, rotting plant debris, bacteria, small invertebrates and bodies of dead tadpoles; in a tadpole population, most are omnivores with small mouthparts; a few though with large mouthparts are still omnivorous, but carnivorous habits are more evident and will swallow their smaller omnivorous brethren whole, making them also cannibalistic; although possesses a skin secretion that acts as an irritant and may sway some predators, predation on these frogs abound from visiting shorebirds, crows, raccoons, snakes, in particular: Heterodon nasicus, Thamnophis marcianus, the Sonoran Desert toad, Incilius alvarius consumes juveniles & adults; predaceous aquatic dytiscid beetle larvae attack tadpoles, while carabid tiger beetles in the subfamily, Cicindelinae will attack newly metamorphosed froglets.

Habits: This frog spends 8-10 months out of the year hibernating in burrows and is well adapted due to this in living in arid environments; dig burrows by backing into ground with hind limbs aided by spades; rocking the body as they dig, soil is thrown upwards falling onto the backs of the body; adults are nocturnal when on land, seeking refuge under rotting logs or plant debris during daylight hours; will become active diurnally during a passing thunderstorm; newly metamorphosed froglets do show diurnal activity until they burrow to

6. S. couchii burrowing

hibernate; has the ability to eat up to 55% of body weight in a single day; once this is done once or twice, it is enough to get the frog through its hibernation period; in hibernating underground; stores nitrogenous wastes to conserve body water; fluid concentration on average is 600 mOsm/l; urea in urine accounts for half that concentration; dilated bladder stores urine which may be utilized to replace body fluid deficits; stores and metabolizes fat; the monogenean, Pseudodiplorchis americanus (a type of flatworm) infects the frog by entering the bladder and feeding off the frogs blood; it would appear that during hibernation, this could be fatal to the frog; but even with heavy infestation of up to 35-95 flatworms per frog, S. couchii may be weak and have great loss of body fat, but survives to end its hibernation and replenish food stocks with no evidence of mortality; the signal of emergence from hibernation are the oncoming signs of spring thunderstorms; are cued by thunder and the low frequency sounds of raindrops; they are described as local migrants as they can migrate up to 200km/124.27mi from hibernation grounds to breeding waters; S. couchii are prolific breeders with all congregating in mass around and in temporary rainwater washes, ephemeral ponds or vernal pools to mate;

7. Tadpoles

though most breeding is timed for first year's thunderstorm, breeding adults are from 2-10 years old; breeding does occur when rain presents itself throughout summer; the males call in water with a plaintive low 'wbaa-wbaa' much like a sheep bleating; most mating and oviposition occurs on first night, once amplectic external fertilization of eggs take place the female lays up to 3,000 eggs adhering them to submerged structures such as plant stalks or sticks; S. couchii eggs & larvae are one of the fastest developing frogs and the fastest in N. America; dependent on temperature, eggs hatch anywhere from 15-36 hours; while tadpoles fully metamorphose into froglets within 7-8 days; the drying of water beds accelerates development, but the quicker developing tadpoles result in smaller sized froglets; the quick mating, egg incubation development and full metamorphosis of larvae are all in response to the drying up of the temporary water bodies; fully metamorphosed juveniles remain by the drying up water for a few days then move to shrub cover until the soil completely dries up where they then burrow into cracks in seeking refuge; they now are nocturnal emerging with the adults at night to forage until they follow adults to permanently burrow and hibernate for the dry season; this species is listed with stable populations and is not threatened.

1. False tomato frog ~ D. guineti

False Tomato Frog
Family: Microhylidae
Genus: Dyscophus
Species: Dyscophus guineti

Length: ♂ ~ 63.5mm/2.5in; ♀ ~ 101.6mm/4in
Weight: ♂ ~ 40.1gm/1.41oz; ♀ ~ 222.55gm/7.85oz
Longevity: 5-10yrs; maximum: 13yrs in wilds
Name Origin: Dyscophus guineti ~ Dyscophus is derived from the Greek word duskophos meaning 'discoid' in reference to flat circular back; guienti honors the French merchant, M. Guinet, who provided the first specimens to science; altogether refers to, 'Guinet's disc-shaped frog'.

Distribution: The false tomato frog is one of 3 tomato frogs belonging to the genus, Dyscophus that all are only found occurring naturally on the island of Madagascar; the 3 are: D. guineti, D. antongilli & D. insularis; further, D. guineti is restricted to patched areas along the eastern rainforests' rim of the island; prefers low lying subtropical-tropical forests with stagnant or very slow moving waters; readily accepts shallow pools and swamp habitat.

2. Note flat dorsum & dorsolateral fold

Description: Most distinguishing feature besides color is the flatness of back and the extended dorsolateral folds on each side; even though this frog belongs to the family; Microhylidae, referring to small narrow mouthed frogs, D. guineti, by no means is little nor possesses a small mouth; microhylid taxonomy is more complicated than that, where morphology presents similarities in short snouts, stout hind limbs, orbicular & teardrop-shaped bodies; skin is primarily smooth and moist; although indistinct, tympanum is one-third the size of eye; set within a green iris, the pupil is slightly horizontally elliptical; the body is short and stocky; toes are long and thin with no webbing on front toes, while hind feet possess very little expressed

3. Note lack of feet webbing

membrane webbing; tibiotarsal articulation reaches tympanum when forelegs are folded; sexual dimorphism is evident in female being larger and brighter in color than male; toes on both sexes usually are tipped white; normally there is a rhomboid dark marking beginning at the back of the eyes extending down the sides below the dorsolateral folds ending at mid-flank; also a wide dark stripe may appear connecting the eye with the mouth; overall conspicuous dorsum & leg coloration varies from shades of orange to reddish brown in some males, while some are yellowish; often there are many small darker reddish orange reticulations; ventrum is white to a creamy white to offish yellow usually with black spotting on throat; juveniles display a bluish gray body and a dorsal dullish tan brown, which becomes brighter as they age; there are no subspecies.

4. Yellow specimen

Diet: False tomato frogs are hearty eaters and will dine primarily on terrestrial arthropods and their larvae such as insects and spiders; are not particularly good swimmers, but occasionally will go after aquatic invertebrates along shoreline; may also take small vertebrates and at times will exhibit cannibalism with small juveniles being taken by adults; juveniles consume smaller invertebrates while tadpole larvae are filter feeders straining any organic material suspended or settled in water; due to body mucous secretions of frog, there are no known habitual predators except for the casual one that hasn't learned yet the lesson of seizing a false tomato frog.

5. juvenile coloration

Habits: This frog is primarily nocturnal; during daytime, usually stays put and concealed excavated down to 6cm/2.4in in ground litter just near a water bodies' edge; at night prefers to walk when foraging, but will hop and jump when need arises; hunting is usually terrestrial, but will also go in shallow water to forage; utilizes ambush strategy; when prey is sighted will hop or lunge toward victim then flick the sticky tongue out to capture prey and bring it back into mouth for swallowing; primarily found in habitat averaging 26.7° C /80° F, but is tolerant of temperatures ranging from 10-35° C/50-95° F from sea level to an elevation of 900m/2,970ft; most restrictive range is humidity, requiring 70-80% humid conditions to keep moist skin from desiccation; the hard & large inner metatarsal tubercule located on underside of each foot protects feet from forest debris and aids in digging; for defense, the frog will stand up on all legs and bloat itself by inhaling air to make it appear larger than actually is; if this bluff doesn't work and is grabbed by a predator, it will secrete a milky & adhering protease peptide from its cheek areas that act as an efficacious irritant to mucous membranes, such as mouth and eyes; peptide is a trypsin inhibitor and most probably had

6. Bloated from air inhalation

originally evolved as an antimicrobial aid to combat invasive microorganisms; predator deterrence was an alternative evolvement; although the proteinase inhibitor found in the frog's skin is not a true poison, once in contact with the sticky substance some humans can experience from mild to serious allergic reactions; the inhibitor congeals once interacting with mucous membranes and in taking days to degrade, will give a careless predator or human days of grief in temporary blindness and blockage of mucous lined orifices such as the mouth & throat; reproduction occurs during cooler months of year; males will first enter stagnant or slow moving shallow waters to begin calling; call is described as a fast rhythm of 'papa-papa-papa;' amplexus is performed in water; as the female releases her eggs, male inseminates; female oviposits between 1,000-1500 eggs on the water's surface; larvae hatch from only a 36 hour incubation period; upon hatching; tadpoles are 0.6cm/0.25in in length upon hatching; within a year tadpoles fully metamorphose into juveniles; maturity for juveniles is reached in 9-14 months; this frog tends to adapt to man's natural habitat destruction and is currently not listed as threatened.

AMPHIBIAN PHOTO CREDITS:

DISCUSSION: 1. George Dewey; 2. illustrator Bob Strauss; 3. top/Jervik, middle/Clack, bottom/Ahlberg; 4. illustrator Nobu Tamura; 5. illustrator Nobu Tamura; 6. sketch artist Alain Beneteau; 7. source: Prehistoric Planet; 8. illustrator George Langford (1876-1964).

SIRENS: 1. Jack Dermid; 2. Laurie Brice (wildlife photographer).
Siren Species: P. axanthus- 1. Todd Pierson (2010); 2. U.S. Geological Survey. S. intermedia texana- 1. Barry Mansell. S. lacertina- 1. D. Byres; 2. source: Bluegrass Aquatics.

CAECILIANS: 1. Ashley Gosselin Ildari; 2. courtesy: Marvalee Wake; 3. illustrator Nobu Tamura.
Caecilian Species: A. eiselti- 1. R. A. Nussbaum. S. thomense- 1. source: California Academy of Sciences; 2. D. Lin (Weckerphoto - GG III); 3. R. A. Nussbaum; 4. John Measey. D. mexicanus- Jonathan Campbell (UTA); 2. Joel Sartare; H. squalostoma- 1. Peter Janzen; 2. Dante Fenolio; 3. Frank Teigler; 4. Dante Fenolio. G. danieli- 1. Varad Giri; 2. Varad Giri. S. annulatus- 1. Andres Acosta; 2. Andres Acosta; 3. M. Wilkinson, compiled by Mark Wilkinson, Alexander Kupfer, Rafael Marques-Porto, Hilary Jenkins, Marta M. Antoniazzi & Carlos Jared. S. kirkii- 1. source: coxi.blog.83fc2.com (Serpent Gothique); 2. source: Alexi Mikailov. U. menoni- 1. Virek Philip Cyriec; 2. Varad Giri. I. kohtaoensis- 1. Nonn Panltuong; 2. Wynaad 2005 (Wikimedia Commons); 3. Marvalee Wake; 4. Nonn Panltuong; 5. source: blog.livedoor.jp (Japan); 6. blog.livedoor.jp (Japan). R. bivittatum- 1.Henrik BrengsØe; 2. source: Natural History Museum (London). T. natans- 1. courtesy: Fort Worth Zoo; 2. Klaus Draeby; 3. Mikalia31 (You Tube).

SALAMANDERS: 1. illustrator Dustdevil; 2. illustrator Michael Skrepnick; 3. Joe Garcia; 4,5&6 source: courses.wahington.edu; 7. source G.L. Guillermo amersol.edu.pe; 8. Gary Mezaros; 9. Stephan M. Deban.

Salamander species: A. means- 1. source: generalexotics.com; 2. D. Dennis; 3. T. Johnson; 4. Barry Mansell; 5. & 6. Margaret Gunzburger; 7. source: blog.livedoor.jp (Japan). N.maculosus- 1. E.R. Degginger; 2. John Cancalosi; 3. Gary Meszaros. C. alleganiensis- 1. Andrew Hoffman; 2. Mike Pinder; 3. Greg Lipps; 4. courtesy: Animaniacs Aquariums & Exotics; 5. courtesy: St. Louis Zoo (Herpetological & Aquatics Dept.) E. rathbuni- 1. Gary Nafis; 2. courtesy of Digimorph staff, University of Texas; 3. Dante Fenolio; 4. Chip Clark; 5. Robert & Linda Mitchell; 6,7,8 courtesy of San Marcos National Fish Hatchery System. P. yonahlossee- 1. Gary Nafis; 2. Timothy Burkhardt; 3. Bill Peterman 4. Wikipedia; 5. source: thehibbits.net; 6. Gary Nafis. B. odonnelli- 1. Sean Michael Rovito; 2. & 3. Todd Pierson. O. pacificensis- 1,2 & 3 Vide Ohlin. N. viridescens- 1. Kevin Stohlgren; 2. George Grall; 3. John M. Coffman; 4. Dylan March; 5. Tom Murray. T. shanjing- John P. Clare; 2. Heather Bjornebo; 3. Henk Wallays; 4. Bill Wergelewski; 5. Henk Wallays. I. alpestris- 1. Frank Deschandol; 2. Henk Wallays; 3. Christoph Riegler; 4. Frank Deschandol; 5. Piet Spaans. A. annulatum- 1. Michael Graziano; 2. Kory Roberts; 3. Bill Peterman; 4. Jason Ksepka. P. anguinus- 1. Arne Hodalic; 2. source: Association of Managers of Slovenia; 3. Rollin Verlinde; 4. Jeroen Speybroeck; 5. Jan van der Voort; D. ensatus- 1. Richard D. Bartlett; 2. Steven Holt; 3. Randomtruth (Flikr); 4. Jens V. Vindum; R. olympicus- 1. Mike Graziano; 2. William Flaxington; 3. Mike Graziano. R. sibricus- 1. Henk Wallays; 2. Dr. Altay Zhatkanbayev; 3. & 4. Henk Wallays; 5. Dr. Altay Zhatkanbayev.

FROGS: 1. source: thalabeach.com.au (Australia); 2. source: batrachos.com (Russia); 3. Kevin Walsh; 4. Todd Marshall; 5. Justin Touchon & Karen Warkentin; 6. Michael & Patricia Fogden; 7. J. Andrews.
Frog species: B. exsul- 1. & 2. Gary Nafis; 3. Dan Suzio; 4. Gary Nafis; 5. Ceal Kingler. R. dorsalis- 1. & 2. Paddy Ryan; 3. Pstevendactylus, source: Wikimedia commons; 4. Scott Wahlberg; 5. Guillermo Lopez Leal. 6. courtesy of Texas Parks & Wildlife Department (TPWD). A. callidryas- 1. Carey James Balboa; 2. Dr. Peter Weish; 3. Jenny de Malaga ; 4. Gerry Marantelli ; 5. Jazu; 6. Robbin Moran; 7. Dr. Peter Janzen. P. terribilis- 1. Devin Edmonds; 2. source: frogpod.blog; 3. source: z00z00.livejournal.com (Flying Ark ZOOZOO); 4. source: Mongabay.com; 5. source: dendrobatenwelt.de (Germany). L. pakeka- 1. Phil Bishop; 2. IUCN Red List/Frogs Australia Network; 3. & 4. Dr. Paddy Ryan; 5. courtesy: Karori Wildlife Sanctuary. C. gayi- 1. Felipe Rabanal; 2. Rune Midtgaard; 3. source: pacmanfrogs.de (Germany); 4. & 5. courtesy: Fundacion Jardin Botanico Nacional. L. fragilis- 1. Esteban Alzate; 2. Isaac Lopez; 3. Andrey Anzofeifa; 4. Kevin Kriger, courtesy of Save the Frogs (STF); 5. Marcos Guerra. P. kl. esculentus- courtesy of Surrey Amphibian and Reptile Group (SARG); 2. Peter Switzer; 3. Anna S. source: Wikipedia; 4. Milan Vogrin; 5. courtesy of SARG. R. catesbeiana- 1. Jim Harding; 2. Dr. Udo M. Savilli; 3. Dwight Kuhn; 4. Richard Seaman; 5. Douglas Alden Peterson; 6. courtesy of U.S. Dept. of Interior/U.S. Geological Survey. H. australiacus- 1. David Nixon; 2. courtesy: New South Wales Government Environment Dept. 3. courtesy: South Coast Herpetological Society Inc.; 4. Ben Brown; 5. Frank Lemckert; 6. Tnarg 12345. A. obstetricans- 1. Daniel Phillips; 2. Jordi B. Colomar; 3. Hardworkinghippie (Flikr); 4. Hugo F. Sousa; 5. Hardworkinghippie (Flikr); 6. Daniel Phillips. C. ilex- 1. Michael Williams; 2. Brad Wilson; 3. Dante Fenolio; 4. Joel Sartore; 5. source: La Taberna del Drunkerosteus. S. couchi- 1. Jack Goldfarb; 2.&3. Gary Nafis; 4. Jim Rorabaugh; 5. Gary Nafis; 6. source: Super Stock; 7. T. Beth Kinsey. D. guineti- 1. Michael Sloviak; 2. source: knowledgeclips.com 3. Francesco Vitali; 4. source: Studio Evenaar (Dutch); 5. David G. Barkasy; 6. Dr. Peter Janzen.

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