61.
Veracruz Pigmy Salamander
(Thorius pennatulus)
CR
Overview
The Veracruz pigmy salamander is a relatively small species, growing to less than 50 mm in total length. This species requires moist environments in which to hide, such as damp rock crevices, rotting logs and leaf litter. Habitat alteration as a result of extensive agricultural activities throughout this species range has reduced the area of suitable habitat, converting much of the original forest into shadeless, open areas. It has experienced more than an 80% of the wild population has been lost in the last 15 years. It has been speculated that the adverse environmental impacts of a volcanic eruption, or even virulent disease (such as chytridiomycosis), might have contributed to the recent dramatic decline.
Urgent Conservation Actions
Surveys to determine species range; disease screening; development of a Conservation Action Plan; disease screening; biodiversity-friendly shade coffee.
Distribution
Central western Veracruz, Mexico
Media from ARKive
ARKive image - Veracruz pygmy salamander
ARKive image - Veracruz pygmy salamander on leaf litter
ARKive image - Veracruz pygmy salamander, dorsal view
ARKive image - Veracruz pygmy salamander curled up on leaf litter
Evolutionary Distinctiveness
Order: Caudata
Family: Plethodontidae
The Plethodontidae is by far the largest family of salamanders, comprising nearly 70% of all living species. In total there are 378 known plethodontids divided between four subfamilies and 24 genera. The plethodontids are united by the fact that they do not possess lungs and breathe entirely through their skin and mouth lining. They are often referred to as the lungless salamanders, although they are thought to have evolved from highly aquatic, lunged ancestors in the streams of the Appalachian Mountains in eastern North America. The earliest plethodontids were hypothesised to have lost their lungs because individuals with reduced, or absent, lungs were less likely to float away in the swift mountain streams where they lived. The vast majority of other salamanders possess lungs, so this makes the lungless salamanders an unusual and fascinating group of animals.

They are thought to have diverged from all other amphibian species 145 million years ago at the boundary between the Jurassic and Cretaceous periods. They are as different from all other amphibian lineages as wombats are from whales, evolving at a time when dinosaurs were still dominant. Overall, plethodontids are the most evolutionary advanced salamanders, so it may at first appear odd that they should have lost lungs, which are one of the most basic features of all vertebrates living on land. Lacking lungs and being dependent on their skin for respiration places a size restriction on these salamanders because large animals have a relatively small surface area of skin compared to their body’s volume, and have greater difficulty in supplying their body tissues with oxygen compared to smaller animals (which have a large surface area to volume ratio). The long, slender form of the lungless salamanders maximises the surface area available for gas exchange, and some species grow to lengths of over 300 mm.

Plethodontid salamanders occupy a great diversity of habitats, ranging from strictly aquatic to strictly terrestrial, exploring niches as diverse as caves, trees, mountain streams, and they are also found burrowing through the earth. Dependence on their skin for breathing places limitations on where and how lungless salamanders can live. Their skin must be kept moist at all times in order for oxygen to be taken up by the blood in capillaries beneath the skin. This means plethodontids are either confined to humid areas, or must find damp hiding places and only emerge in wet weather, typically at night. The life of a lungless salamander in less humid areas, like Europe and temperate North America, therefore comprises brief periods of activity interspersed with inactive phases that are often very long. They are able to survive the periods of inactivity because they have a very low metabolic rate and low energy requirements. Able to store much of what they eat as fat, they do no need to feed very often.

A further adaptation, present among many species of the lungless salamander subfamilies named “Plethodontinae” (from East and West North America) and “Bolitoglossinae” (from tropical Central and South America), is “direct development”. This is a method of amphibian development where the larval stage (e.g. the tadpole stage in a frog’s life history) has been eliminated. Early development takes place in eggs, which may be laid in moist places away from water, and the young hatch out as miniature adults. The well known amphibian metamorphosis, most commonly appreciated in the transition from tadpole to adult frog, does not occur outside of the egg. This mean that certain lungless salamanders in these two subfamilies may live away from water bodies, allowing them to expand their ranges to new areas.

The history and characteristics of the lungless salamanders go some way to explaining their range. They are mostly found in the New World, where they are widely distributed in eastern and western North America, as well as Central and South America. However, continental drift over millions of years has also brought them to the Old World, where they are found in parts Europe (e.g. Sardinia) and Korea. The existence of the Korean crevice salamander was unknown until 2005, when its discovery was a shock to science, indicating a long history of lungless salamanders in Asia. This is the only known species is Asia, suggesting that the rate of species generation in this part of the world is very low, especially compared the the huge radiation of lungless salamander species in the New World.

There are 23 known species in the genus Thorius (commonly known as the “Mexican pigmy salamanders”) which represent one of ten genera present within a lungless salamander subfamily called the “Bolitoglossinae”, including all the plethodontids from Central and South America. The Mexican pigmy salamanders include the smallest salamanders on earth – the most diminutive of which measure just 26.9 mm in total length as sexually mature adults. Thorius salamanders occur only in Mexico, where they are restricted to the southern states of Veracruz, Puebla, Oaxaca and Guerrero. In general, Mexican pygmy salamanders live at high elevations, ranging from about 1,500-3,000 metres above sea level or higher, but some species descend to 800 metres above sea level.

Most Mexican pigmy salamanders lack teeth and their skulls are extraordinary because of the poor state of development, the thinness of the bones, and the weak articulation of the elements. Miniaturisation has been achieved by the reduction or loss of some of the cranial (or skull) elements, accompanied by a relative increase in the size of the sense organs. Another interesting feature of these salamanders is that they possess male heterogamy reproduction – the presence of an X or Y-type sex chromosomes in the eggs and sperm, as is the case in humans. This is known as chromosomal sex determination, where females have two X sex chromosomes (XX) in their cells and males have one X chromosome and one “male” Y chromosomes (XY). In non-chromosomal sex determination, being male or female can occur as a result of environmental conditions, such as temperature, whereas with the X and Y-chromosome system, sex is determined from the outset.

The genus Thorius is thought to have originated in the Early to Mid Miocene period, between 23 and ~12 million years ago. This makes Mexican pigmy salamanders as dissimilar to their closest relative as humans are to gibbons. Specifically, within the Thorius salamanders the Veracruz pigmy salamander diverged about 5 million years ago, and is closely related to another top 100 EDGE species: the San Martìn Pigmy salamander (Thorius narismagnus).
Description
The Veracruz pigmy salamander, like all lungless salamanders in the Bolitoglossinae subfamily, possesses a slender body, long tail and prominent eyes. A distinctive feature of the plethodontid family is a narrow groove (the nasolabial groove) running from each nostril to the upper lip: its function is to carry waterborne odours from the ground into the nasal cavity. Another curious trait of the lungless salamanders are mental (from the Latin “mentum”, meaning chin) glands. These are modified mucus glands and release pheromones, which are chemicals produced by an animal to influence the behaviour of other members if its species, often with regard to breeding receptivity. Mental glands are sometimes visible in males as raised bumps below their lower lip.

Lungless salamanders are very small to medium in size, usually measuring between 25 to 250 mm from the tip of the nose to the end of the tail, which salamanders retain throughout their life. They are unusual among the salamanders in that some species can detach from their tail as a predator-defence mechanism (also known as tail or caudal autotomy). It is therefore not unusual to see individuls missing part or all of their tail, which they may regererate later. Lungless salamanders may have bold patterns on their skin as adults, or they may have a colouration more similar to their environment to aid camouflage. They have well-developed “costal” grooves (successive vertical grooves in the skin along the sides of the body), generally numbering between 12-15. Their limbs are slender and often have largely or completely webbed digits. Species, like the Veracruz pigmy salamander, in the Thorius genus (the “Mexican pigmy salamander”) are very similar in form to those in the genus Bolitoglossa (the “mushroom-tongue salamanders”), although they are generally much small, the largest representatives seldom exceeding 70 mm in total length.

The Veracruz pigmy salamander is a small species, growing to a total length of 49 mm, 23 mm of this measurement being accounted for by the stout tail. The head is slightly wider than the neck and the snout projects slightly beyond the mouth. There are 13 “costal” grooves along either side of the body. This species has slender, weak limbs with very small digits. The skin is smooth, and yellowish brown in colour along the dorsal (or upper) surface. A grey band extends along the back to the tip of the tail. This band is particularly light and slightly metallic along its undulating or serrated margins, and is marked by a number of short V-shaped spots which appear in pairs, one to each costal band (the body segments bordered by the costal grooves). The colouration of the ventral surface (or underside) is brown.
Ecology
Most Mexican pigmy salamanders are usually terrestrial (or ground-dwelling), living mainly under surface cover, inside logs, or especially beneath the bark of fallen and rotting logs. In 1908, the German naturalist Hans Friedrich Gadow in his publication “Through southern Mexico, being an account of the travels of a naturalist” remarked after observing some members of this family:

These little things showed a predilection for living in a proverbially precarious position, namely, “between the bark and the wood” of decaying pine-trees, amongst the boring-dust of beetles and maggots.

Very little is known about the Veracruz pigmy salamander, although it thought to be ground dwelling and breeds by direct development, with the young maturing within the eggs and hatching as miniature adults. This whole process is independent of a water body, making this a truly land-dwelling species. This species is ground dwelling and requires moist conditions, such as those found in rock crevices, under rocks, and in leaf litter, as well as within and under rotting logs. It also occurs in shade coffee plantations where humidity is maintained Some species in the Thorius genus are known to display courtship rituals. The pheromone releasing mental gland on the chin of male Mexican pigmy salamanders plays an important role in mating to influence the receptivity of females. During amplexus (the mating embrace), the male clasps the female with both his arms and legs, and rubs pheromones across the female’s snout. Female Mexican pigmy salamanders have been found to guard the eggs throughout their development in many species, often in special hides, until hatching occurs.

Mexican pigmy salamanders may at first appear very vulnerable to predators but a number of defense mechanisms have been found among the members of this genus. These include caudal autotomy (tail detachment) and behavioural defensive methods, including immobile posture, coiling and flipping of the body, and displays where the stomach is exposed and the tail is held up or undulated.
Habitat
The Veracruz pigmy salamander inhabits cloud forest, and also forest at lower altitudes. It requires moist situations such as within rock crevices, under rocks, and in leaf litter especially at the base of banana trees), in addition to in and under rotting logs. This species can tolerate a certain amount of habitat alteration in that it has been found to occur in shade coffee plantations where a high humidity is maintained.
Distribution
This species is known only from central western Veracruz, from Soledad Atzompa in the south and west, east to Cerro Chicahuaxtla, north to Teocel, Mexico, generally at altitudes of 1,000-1,200 metres above sea level (although it has been found as high as 2,000 metres).
Population Estimate
Although once very common, this species appears to have undergone a dramatic decline of over 80% in the past 15 years. The Veracruz pigmy salamander started to disappear in the 1980s. Since then there have been very few sightings of this species. There was one record in the late 1990s and a single individual was observed in 2004.
Population Trend
The IUCN Red List of Threatened Species indicates that the total population size of the Oaxacan pigmy salamander is in decline.
Status
The Veracruz pigmy salamander is listed as Critically Endangered in the IUCN Red List of Threatened Species because of a drastic population decline, estimated to be more than 80% over the last three generations, inferred from the apparent disappearance of most of the population. The generation length is assumed to be five years. In addition, this species is protected by Mexican Law under the "Special Protection" (Pr) category.
Threats
The main threat to the Veracruz pigmy salamander is the transformation of extensive areas of its forest habitat into open unshaded agricultural habitats. Since this species requires a high degree of humidity, it cannot survive where habitat alteration has brought overly dry conditions, not least because lungless salamanders must maintain moist skin to enable them to breathe. The forests have been severely reduced by logging, as well as by expanding human settlements as the area becomes increasingly urbanised. However, these threats do not explain the level of decline that has been observed, since the habitat is still in quite good condition in a few places. It is speculated that the adverse environmental impacts of a volcanic eruption, or virulent disease (such as chytridiomycosis, although this normally impacts species that are associated with water), might have caused this drastic decline.
Conservation Underway
This species has not been found in any protected areas, although it is protected by Mexican Law under the category "Special Protection" (Pr).
Projects

  • Establish spatial knowledge of the distribution and occurrence of some of the threatened species in Mexico.
  • Screen individuals and other amphibians in these areas for the presence of chytrid fungus.
  • Increase the capacity in Mexico for the long term conservation of these species

Conservation Proposed
Habitat protection is urgently required to maintain hogh quality habitat for this species. Furthermore, suitable closed and humid habitat should be maintained. The Veracruz pigmy salamander has been found to survive in shade-grown coffee plantations so this could be a land use encouraged throughout the range of this species, in addition to any other more biodiversity friendly agricultural practices. Research is also needed to establish the reasons for this species’ population crash, including disease screening and a thorough examination of the factors that are thought to threaten the Veracruz pigmy salamander. Additional surveys are also required to determine exactly where (if at all) populations survive in the wild. All of the resultant information should be used to create a Conservation Action Plan as a vital first step in galvanising efforts to preserve this species should it still survive in the wild.

If the species is found to survive in the wild, it is essential that its habitat is protected to prevent further population extinctions. In addition to conserving any remaining wild habitat for this species, the IUCN Technical Guidelines for the Management of Ex situ Populations, part of the IUCN Red List of Threatened Species, recommend that all Critically Endangered species should have an ex situ population managed to guard against the extinction of the species. An ex situ population is ideally a breeding colony of a species maintained outside of its natural habitat, giving rise to individuals from that species that are sheltered from problems associated with their situation in the wild. This can be located in the specie’s range or in a foreign country that has the facilities to support a captive breeding programme for that species. Since the Upper Cerro pigmy salamander is currently categorised as Critically Endangered, the possibility of a captive breeding programme for this species should be investigated if future surveys discover any remaining populations.
Associated EDGE Community members

James is an expert in evolutionary morphology, developmental biology, and systematics.

Links
References
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Dunn, E. R. 1926. The salamanders of the family Plethodontidae. Smith College, Northampton, Massachusetts, U.S.A.

Frost, Darrel R. 2006. Amphibian Species of the World: an Online Reference. Version 4 (17 August 2006). Electronic Database accessible at: . American Museum of Natural History, New York, USA.

Frost, D. R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C. F. B., De Sá, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M., and Wheeler, W.C. 2006. The Amphibian Tree of Life. Bulletin of the American Museum of Natural History 297: 1-370.

Gadow, H. 1908. Through southern Mexico, being an account of the travels of a naturalist. Witherby and Co., London.

Halliday, T. and Adler, C. (eds.). 2002. The new encyclopedia of reptiles and amphibians. Oxford University Press, Oxford.

Hanken, J. and Wake, D.B. 1998. Biology of tiny animals: systematics of the minute salamanders (Thorius: Plethodontidae) from Veracruz and Puebla, Mexico, with descriptions of five new species. Copeia 1998: 312-345.

IUCN, Conservation International and NatureServe. 2006. Global Amphibian Assessment. Global Amphibian Assessment. Accessed on 08 December 2006.

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Macey, J. R. 2005. Plethodontid salamander mitochondrial genomics: A parsimony evaluation of character conflict and implications for historical biogeography. Cladistics 21:194-202.

Min, M. S., Yang, S.Y., Bonett, R.M., Vieites, D. R., Brandon, R.A. and Wake, D.B. 2005. Discovery of the first Asian plethodontid salamander. Nature 435:87-90.

Mueller, R. L., Macey, J.R., Jaekel, M., Wake, D.B. and Boore, J.L. 2004. Morphological homoplasy, life history evolution, and historical biogeography of plethodontid salamanders inferred from complete mitochondrial genomes. PNAS 101:13820-13825.

Obst, F.J., Richter, K. and Jacob, U. 1984. The Completely Illustrated Atlas of Reptiles and Amphibians for the Terrarium. T.F.H. Publication Inc., N.J., U.S.A.

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Parra-Olea, G., García-París, M. and Wake, D.B. 1999. Status of some populations of Mexican salamanders. Revista de Biologia Tropical 47: 217-223.

Parra-Olea, G., Papenfuss, T.J. and Wake, D.B. 2001. New species of lungless salamanders of the genus Pseudoeurycea (Amphibia: Caudata: Plethodontidae) from Veracruz, Mexico. Scientific papers of the Natural History Museum of the University of Kansas 20: 1-9.

Roelants, K., Gower, D. J., Wilkinson, M., Loader, S. P., Biju, S. D., Guillaume, K., Moiau, L. and Bossuyt, F. 2007. Global patterns of diversification in the history of modern amphibians. Proceedings of the National Academy of Sciences 104: 887-892.

Sever, D.M. (ed.). 2003. Reproductive biology and phylogeny of the Urodela. Science Publishers, Inc., New Hampshire, U.S.A.

Wake, D. B. 1966. Comparative osteology and evolution of the lungless salamanders, family Plethodontidae. Memoirs of the Southern California Academy of Sciences 4:1-111.

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