90.
Smith's False Brook Salamander
(Pseudoeurycea smithi)
CR
Overview
Smith’s false brook salamander used to be very abundant; there was a time when hundreds could be seen in a day. However, it is now extremely rare, suggesting that a massive population decline has taken place. The last living specimen may have been seen by Dr. David Wake (expert in the Plethodontidae) in the late 1990s. For the last 15 years it has been almost impossible to find. The Smith’s false brook salamander is thought to be closely related to another top 100 EDGE plethodontid: the aquatic false brook salamander, which is the only known aquatic lungless salamander in between Northern Mexico and Panama, all the other species being ground dwelling.
Urgent Conservation Actions
Strengthen the protection of remaining habitat and determine the reasons for its dramatic decline.
Distribution
North-western Oaxaca, Mexico.
Fact
Smith’s false brook salamander was named in honour of Dr. Hobart M. Smith who discovered the species in 1935.

Smith’s false brook salamander is thought to be closely related to another top-100 EDGE amphibian that occurs near to its range: the possibly extinct aquatic false brook salamander, previously found in the northern mountain slopes of central-eastern Oaxaca, Mexico.
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, making 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 evolutionarily advanced salamanders, so it may at first appear odd that they should have lost their 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 not 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 means 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 of 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 in Asia, suggesting that the rate of species generation in this part of the world is very low, especially compared to the huge radiation of lungless salamander species in the New World.

Comprising 50 known species, the genus Pseudoeurycea (commonly known as the “false brook salamanders”) is one of the largest genera in the Plethodontidae family, second only to the Bolitoglossa genus (the “mushroomtongue salamanders” – 93 known species). The false brook salamanders are present within the subfamily “Bolitoglossinae” (from Central and South America) and its members are very wide ranging in size, with some of the smallest and largest lungless salamanders included (total lengths from about 65 mm to 325 mm).

The false brook salamanders have recently been reorganised and expanded to absorb a number of other genera that are now considered to fit within the Pseudoeurycea genus. The taxonomy of these salamanders may not be finally resolved just yet, but it is known that the whole group diverged from all other salamanders in the Late Eocene period, at least 34 million years ago. This is around the same time that humans and monkeys shared a common ancestor.

The Smith’s false brook salamander is thought to be closely related to another top 100 EDGE plethodontid: the aquatic false brook salamander, the only known aquatic lungless salamander in between Northern Mexico and Panama, all the other species being terrestrial (or ground dwelling).
Description
Smith’s false brook 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 (chemicals produced by an animal to influence the behaviour of other members of 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 regenerate 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 Smith’s false brook salamander, in the genus Pseudoeurycea (the “false brook salamanders”) are very similar in form to those in the genera Chiropterotriton (the “splayfoot salamanders”) and Bolitoglossa (the “mushroom-tongue salamanders”).

Smith’s false brook salamander is a fairly large species, reaching total lengths of about 145 mm, about 76 mm being accounted for by the proportionally long tail. The tail is constricted at the base and is slightly thicker and shorter in males than in females. The head is broad with strongly swollen parietal regions (the glandular areas either side of the head behind the eyes) and males possess a distinct mental gland. Thirteen “costal” grooves are present in the skin along either side of the body. The limbs are well-developed and the digits are partially webbed. The dorsal (or upper) surfaces of this species are dark brown, with the sides of the body being yellowish or greyish olive-brown tinged with red. A series of 13 vertical blackish bars are found on the sides of the body. The colouration of the chin is cream with a peppering of cinnamon-brown spots. The belly is greyish-cream with traces of darker lines following the costal grooves. There are numerous blackish spots along the sides of the tail.
Ecology
Like other false brook salamanders, Smith’s false brook salamander is probably active all year round and mating is thought to occur throughout the year. Some species in this genus are known to display courtship rituals. The pheromone releasing mental gland in males 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. Fertilisation is internal and eggs will generally be laid at the beginning of the dry season in November. Female false brook salamanders have been found to guard the eggs throughout their development in many species, often in special hides, until hatching occurs at the beginning of the rainy season. Very little is known about the Smith’s false brook salamander but it is understood that direct development of the young occurs within the eggs and they hatch as miniature adults. This whole process is independent of a water body, making this a truly terrestrial (or land-dwelling) species.

False brook salamanders are well adapted for moving around their habitat and have prehensile tails, meaning they can use their flexible tail as a fifth limb to grip with and hang from. This may prove particularly useful in arboreal (or tree-dwelling) species, although Smith’s false brook salamander would also benefit from having a strong, flexible tail in its own terrestrial (or ground dwelling) life. False brook 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: noxious skin secretions; a poisonous gland on the back of the head (parotoid glands); warning colours on the skin of the back (also termed aposematic colouration) or camouflage colouration; and caudal autotomy (tail detachment). Behaviourally, false brook salamanders have been found to orchestrate many defensive methods, including immobile posture, coiling and flipping of the body, tail undulation displays and biting.
Habitat
Smith’s false brook salamander inhabits pine forests and is terrestrial (or ground dwelling), living under the bark of logs. It is able to persist in somewhat disturbed habitat.
Distribution
Occurs only on the Sierra de Juarez, Cerro San Felipe and Sierra de Quatro Venados, north-western Oaxaca, Mexico, at an altitude of 2,500-3,000 metres above sea level.
Population Estimate
It used to be very abundant and there was a time when you could see hundreds in a day. However, it is now extremely rare suggesting that a massive decline has taken place. The last living specimen may have been seen by Dr. David Wake (expert in the Plethodontidae) in the late 1990s. For the last 15 years it has been almost impossible to find.
Population Trend
The IUCN Red List of Threatened Species indicates that the Smith’s false brook salamander population is experiencing a severe decline.
Status
Listed as Critically Endangered on the IUCN Red List of Threatened Species because of a large population decline (estimated to be more than 80% over the last three generations, inferred from the apparent disappearance of most of the population), shrinkage in distribution and habitat destruction and degradation in northwestern Oaxaca, Mexico. The generation length is assumed to be five years.
Threats
This species has probably been negatively impacted by the expansion of agriculture, human settlements and logging. However, these threats do not explain the level of decline that has been observed, since the habitat remains in quite good condition in some places. It has been speculated that the adverse environmental impacts of a volcanic eruption might be the cause of the decline. Other possible explanations could be climate change and disease (possibly chytridiomycosis, although this fungal disease normally impacts species that are associated with water).
Conservation Underway
The species is known to occur in the Parque Nacional Benito Juarez and is listed as "Threatened" (Amenazada) by the Mexican government.
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
Continued and strengthened protection of the species' remaining habitat is required, in addition to research into the reasons for its dramatic decline.

In addition to conserving 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 within the species’ range or in a foreign country that has the facilities to support a captive breeding programme for that species. Further investigation is therefore required into the possibility of establishing a captive breeding programme for any surviving Smith’s false brook salamanders. Captive animals could then be a source of new individuals to repopulate any restored habitat.
Links
References
AmphibiaWeb: Information on amphibian biology and conservation [web application]. 2006. Berkeley, California: AmphibiaWeb. Available: amphibiaweb. Accessed: 08 December 2006.

Chippindale, P. T., Bonett, R.M., Baldwin, A.S. and Wiens, J.J. 2004. Phylogenetic evidence for a major reversal of life-history evolution in plethodontid salamanders. Evolution 58:2809-2822.

Duellman, W. E. and Trueb, L. 1986. Biology of Amphibians. McGraw-Hill, New York.

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

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.

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.

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

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

Larson, A. 1991. A molecular perspective on the evolutionary relationships of the salamander families. Evolutionary Biology 25:211-277.

Larson, A. and Dimmick, W.W. 1993. Phylogenetic relationships of the salamander families: A analysis of congruence among morphological and molecular characters. Herpetological Monographs 7:77-93.

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.

Parra Olea, G. & Wake, D. 2004. Pseudoeurycea smithi. In: IUCN 2006. 2006 IUCN Red List of Threatened Species. IUCN Red List of Threatened Species. Downloaded on 11 December 2006.

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.

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.

Taylor, E. H. 1939. Mexican salamanders. The University of Kansas Science Bulletin 25: 269-272.

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.

Wake, D. B. and Larson, A. 1987. Multidimensional analysis of an evolving lineage. Science 238:42-48.

Wake, D. B. and Lynch. J.F. 1976. The distribution, ecology and evolutionary history of plethodontid salamanders in tropical America. Natural History Museum of Los Angeles County Science Bulletin 25:1-65.

Wake, D.B. 1987. Adaptive radiation of salamanders in Middle American cloud forests. Ann. Missouri Bot. Gard. 74: 242-264.

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