44.
Monzón's Moss Salamander
(Cryptotriton monzoni)
CR
Overview
Very little is known about Monzón’s moss salamander because it was only collected once in the late 1990s. It lives in lower montane moist forest in cloud forest conditions, where it has been found living in bromeliads. The major threat to this species is habitat loss around La Unión as a result of extensive logging and agricultural encroachment. It is presumed to be a very rare.
Urgent Conservation Actions
Surveys to establish current population status; development of Conservation Action Plan; protection of remaining habitat.
Distribution
La Unión, Zacapa in Guatemala
Fact
Monzón’s moss salamander was first described in 1998 by Jonathan Campbell and Eric Smith as Nototriton monzoni. For this reason, its common name “moss salamander” still reflects its initial description, although this is the name given to species within the genus Nototriton despite the fact that Monzón’s moss salamander is now placed in the genus Cryptotriton, which was introduced to plethodontid taxonomy in 2000 by Mario García-París and David B. Wake. This species was named in honour of José Monzón, who assisted Jonathan Campbell and Eric Smith during their research in Guatemala.
Media from ARKive
ARKive image - <i>Cryptotriton monzoni</i> crawling across leaf
ARKive image - <i>Cryptotriton monzoni</i>
ARKive image - <i>Cryptotriton monzoni</i>
ARKive image - <i>Cryptotriton monzoni</i> crawling
ARKive image - <i>Cryptotriton monzoni</i>, side view
ARKive image - <i>Cryptotriton monzoni</i>, anterior view
ARKive image - <i>Cryptotriton monzoni</i> on moss-covered tree trunk
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 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.

Cryptotriton is genus comprising just six species, and is one of twelve genera present within a lungless salamander subfamily called the “Bolitoglossinae”, including all the plethodontids from Central and South America. The Cryptotriton genus was only defined as a separate group in 2000, and previously its 6 constituent species had been placed in the genus Nototriton (the “moss salamanders”). However, the evolution of the Cryptotriton genus is thought to predate that of Nototriton. The moss salamanders originated as far back as the early Miocene (around 23 millions years ago), where as the Cryptotriton genus is believed to have arisen even earlier, possibly in the Oligocene period which began 34 million years ago, when the ancestors of humans and monkeys diverged.

The Cryptotriton salamanders are in the same clade (or sub-section of the evolutionary tree of life) as other top 100 EDGE amphibian genera, including Dendrotriton (the “bromeliad salamanders”), Oedipina (the “worm salamanders”) and, of course, the moss salamanders (Nototriton). 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
Monzón’s moss 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. Monzón’s moss salamander, the only species the genus Cryptotriton, is externally similar to species in the genera Nototriton (the true “moss salamanders”) and Chiropterotriton (the “splayfoot salamanders”).

Monzón’s moss salamander is a small species of lungless salamander, with a total length of around 45 mm (of which around 23 mm is accounted for by a fairly lengthy tail). This species has a broadly rounded snout, large nostrils and slightly protruding, copper-coloured eyes. The body is narrow with 12 “costal” grooves found along the sides in the region of the ribs and abdomen. The limbs are relatively long and the hands and feet each have four digits, which are webbed, especially in the case of the feet. The dorsal (or upward facing) surface of the salamander is brown, with small bluish flecks that also appear on the sides of the head and body. Rust coloured streaks appear on the top of the head, the back and the tail. The ventral (or lower) surface of this species is dark drown.
Ecology
The Cryptotriton salamanders, being closely related to the moss salamanders, worm salamanders and bromeliad salamanders, share certain ecological traits with these groups. They are found in Central America and are partially terrestrial (ground dwelling) and also arboreal (tree-dwelling), where they can be found in bromeliads. Bromeliads are a common type of plant from the Neotropics which possess overlapping leaves that may trap water and organic debris, providing a habitat for numerous species of invertebrate, amphibian and other creatures. They are often found growing on trees, attaching by structural roots to branches or bark.

Very little is known about Monzón’s moss salamander and the Cryptotriton genus in general, although a certain amount of information could be inferable from what is known about their close relatives. For example, although lungless salamanders are generally associated with some degree of parental care through the guarding of their eggs by the female, this feature is lacking in both moss salamanders and worm salamanders. Species within the Cryptotriton genus would need to be studied further to understand whether they do display parental care, but this does not seem to be something that has evolved or been retained throughout their clade.

Direct development of the young occurs 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. Some species in this genus may display courtship behaviour. The pheromone releasing mental gland on the chin of male Cryptotriton 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.
Habitat
It lives in lower montane moist forest in cloud forest conditions, in an area where several isolated mountain peaks arise from relatively dry lowlands. The broad leaf forest habitat contains numerous sweetgums (Liquidambar) and most of the larger trees are heavily laden with epiphytes (plants that grow on other pants, especially trees). The first individual of this species collected for science was found in a bromeliad in primary forest three metres above the forest floor.
Distribution
This species is known only from the original specimen collected, which was found near La Unión, Zacapa in Guatemala, on slopes drained by the Río La Jigua, which flows into the Río Motagua. The only know individual was collected at an altitude of 1,570 metres above sea level.
Population Estimate
No information is available on population status of Monzón’s moss salamander because it has only been collected once in 1998, but it is presumed to be rare.
Population Trend
The IUCN Red List of Threatened Species indicates that the total population size of Monzón’s moss salamander is in decline.
Status
Monzón’s moss 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., all individuals are in a single location, and there is continuing decline in the extent and quality of its habitat in Zacapa, Guatemala.
Threats
The major threat to this species is habitat loss around La Unión as a result of extensive logging and agricultural encroachment.
Conservation Underway
This species is not known from any protected areas, and there are currently no conservation measures underway for this species.
Conservation Proposed
Urgent surveys are needed to establish the current population status of this species and all information collected should then form the basis of a Conservation Action Plan. Protection of the remaining habitat for Monzón’s moss 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 Monzón’s moss salamander. Captive animals could then be a source of new individuals to repopulate protected habitat.
Links
References
Acevedo, M. & Wake, D. 2004. Cryptotriton monzoni. 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).

Campbell, J.A. and Smith, E.N. 1998. New species of Nototriton (Caudata: Plethodontidae) from eastern Guatemala. Scientific Papers of the Natural History Museum of the University of Guatemala 6: 1-8.

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.

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.

García-París, M. and Wake, D.B. 2000. Molecular phylogenetic analysis of relationships of the tropical salamander genera Oedipina and Nototriton, with descriptions of a new genus and three new species. Copeia 2000: 42-70.

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. . 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.

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.

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