44.
Cuchumatanas Bromeliad Salamander
(Dendrotriton cuchumatanus)
CR
Overview
This species inhabits high-elevation, very humid oak forest, where individuals have been found under moss and bark on large fallen trees. It is threatened by habitat destruction, and extensive forest clearance for timber around the only known locality of this species has left only a small remnant patch of oak forest. It is not known for certain whether this species survives in the wild.
Urgent Conservation Actions
Surveys to establish current population status; development of a Conservation Action Plan; protection of remaining habitat.
Distribution
Sierra de los Cuchumatanes, Guatemala
Fact
The genus name for the Cuchumatanas bromeliad salamander, Dendrotriton, is from the Greek “dendron” meaning tree. This is a reference to the arboreal (or tree-dwelling) lifestyle of this group of salamanders. This species is named after the Sierra de los Cuchumatanes in Guatemala where it is found.
Media from ARKive
ARKive image - Forest bromeliad salamander
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 upon 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 upon 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 six species in the genus Dendrotriton (commonly known as the “bromeliad salamanders”), which represent one of twelve genera present within a lungless salamander subfamily called the “Bolitoglossinae”, including all the plethodontids from Central and South America. Although formerly included in genus Chiropterotriton (the “splayfoot salamanders”), the bromeliad salamanders where assigned their own group in 1983 and current evidence suggests that they evolved at least 12 million years ago. This is well over 11 millions years before the origin of modern humans.

The bromeliad salamanders are in the same clade (or sub-section of the evolutionary tree of life) as other top 100 EDGE amphibian genera, including Cryptotriton, Oedipina (the “worm salamanders”) and Nototriton (the moss salamanders). An interesting feature of all of these related genera 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.
Description
The Cuchumatanas bromeliad 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. During amplexus (the mating embrace), the male clasps the female (with both his arms and legs) and rubs pheromones across the female’s snout. 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 10-20. Their limbs are slender and often have largely or completely webbed digits. Species, like the Cuchumatanas bromeliad salamander, in the genus Dendrotriton (the “bromeliad salamanders”) are externally very similar to members of the genus Chiropterotriton (the “splayfoot salamanders”).

The Cuchumatanas bromeliad salamander is small (no more than 80-100mm in total length) and slender with a long, tapering tail. It has long limbs and broad hands and feet, with wide-tipped digits. The head is no wider than the rest of the body and has large protruding eyes and a broad, flattened snout. Its dorsal (or upper) surface is light brown in colour with a darker brown band running down the center of the head, back and tail. The ventral (or underside) colouration is separated from the dorsal colour by a dark drown band along the side of the body. The ventral surface is a greenish brown shade with yellowish blotching.
Ecology
The common family name for arboreal (tree-dwelling) species in the genus Dendrotriton (the “bromeliad salamanders”) is derived from the fact that these salamanders are often found to inhabit a group of plants in the Neotropics called the bromeliads. Bromeliads possess overlapping leaves that may trap water and organic debris, providing a habitat for numerous species of invertebrate, amphibian and other creatures. They often occur growing on trees, attaching by structural roots to branches or bark. These bromeliads are referred to as “epiphytic”, from the Greek “epi” meaning upon and “phyton” meaning plant. The larger tank-forming bromeliads contain a number of substantial pools of water (or small aquaria) between their leaves, and therefore constitute a particularly humid micro-environment within the forest for these lungless salamanders, which are known to favour moist conditions to keep their skin hydrated for better respiration. Oddly enough, however, the Cuchumatanas bromeliad salamander has never been found in bromeliads, but instead under moss and bark on large fallen trees.

The Cuchumatanas bromeliad salamander is a little-studied species, although direct development of the young is known to occur within the eggs and they hatch as miniature adults. This whole process is independent of a water body since the eggs are laid in damp locations on the land, making this a truly terrestrial (or land-dwelling) species. Bromeliad salamanders have been observed displaying courtship rituals. The pheromone releasing mental gland on the chin of male bromeliad 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 bromeliad salamanders may guard the eggs throughout their development in special hides until hatching occurs. However, this behaviour is not found throughout the clade they share with the moss salamanders and worm salamanders – in both of these genera parental care is not observed.
Habitat
This species inhabits high-elevation, very humid oak forest. Individuals have been found under moss and bark on large fallen trees. It has also been found within moss, but none has so far been discovered within bromeliads.
Distribution
This species is only known from a single site south-west of San Juan Ixcoy, in the Sierra de los Cuchumatanes, Guatemala, at an altitude of about 2,860 metres above sea level. However, there is also a probable recent record from Villa Alicia San Martin Cuchumatanes in the same area.
Population Estimate
There is no information on current population status of this species.
Population Trend
The IUCN Red List of Threatened Species indicates that the total population size of the Cuchumatanas bromeliad salamander is in decline.
Status
The Cuchumatanas bromeliad salamander is listed as Critically Endangered in the IUCN Red List of Threatened Species because its extent of occurrence is less than 100 km sq., its distribution is severely fragmented, and there is continuing decline in the extent and quality of its habitat in the Sierra de los Cuchumatanes, Guatemala.
Threats
Although there has been extensive forest clearance for timber around the known locality of the Cuchumatanas bromeliad salamander, a small patch of oak forest is known to remain in the area.
Conservation Underway
The Cuchumatanas bromeliad salamander was originally found inside the proposed Parque Nacional Cuchumatanes, but it is not known whether the species still survives in this area. There are currently no conservation measures underway for this species.
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
Further survey work is needed to determine whether this species survives in the wild, and all information collected should form the basis of a Conservation Action Plan. Protection of the remaining small patch of suitable habitat for the Cuchumatanas bromeliad salamander is also a major priority.

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 possibilities of establishing a captive breeding programme for the Cuchumatanas bromeliad salamander. Captive animals could then be a source of new individuals to repopulate protected habitat.
Links
References
Acevedo, M. & Wake, D. 2004. Dendrotriton cuchumatanus. In: IUCN 2006. 2006 IUCN Red List of Threatened Species. IUCN Red List of Threatened Species.. Downloaded on 11 December 2006.

AmphibiaWeb: Information on amphibian biology and conservation [web application]. 2006. Berkeley, California: AmphibiaWeb. Available: amphibiaweb. Accessed: 08 December 2006.

Campbell, J.A. 2001. Guide to the Reptiles and Amphibians of Guatemala. University of Texas, Arlington. (Web published: http://www.uta.edu/biology/campbell).

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.

Collins-Rainboth, A. and Buth, D.G. 1990. A reevaluation of the systematic relationships among species of the genus Dendrotriton (Caudata: Plethodontidae). Copeia 1990(4): 955-960.

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

Elias, P. 1984. Salamanders of the northwestern highlands of Guatemala. Natural History Museum of Los Angeles County - Contributions in Science 348: 1-20.

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.

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.

Lynch, J.F. and Wake, D.B. 1975. Systematics of the Chiropterotriton bromeliacia group (Amphibia: Caudata), with description of two new species from Guatemala. Natural History Museum of Los Angeles County - Contributions in Science 265: 1-45.

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.

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.

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.

Wake, D.B. and Elias, P. 1983. New genera and a new species of Central American salamanders, with a review of the tropical genera (Amphibia, Caudata, Plethodontidae). Natural History Museum of Los Angeles County - Contributions in Science 345: 1-19.

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