Paghman Mountain Salamander
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Order: Caudata
Family: Hynobiidae
The Hynobiidae family comprises just 50 species, and is commonly referred to as the “Asiatic salamanders”. They are distributed primarily across Asia, although one species is found in European Russia. The hynobiids are an early branch of salamanders and are fairly closely related to the Cryptobranchidae family (the giant salamanders), with which they form the caudate (or salamander) suborder “Cryptobranchoidea”. The earliest fossil record of hynobiids is in the late Miocene of Europe (around 5 million years ago), although recent analyses have indicted that they originated in the Middle Cretaceous about 110 million years ago, at a time when the present-day North American continent was still partly connected to the Europe during the break up of an ancient super-continent called Laurasia. This makes Asiatic salamanders more different from their closest living relatives than humans are to hedgehogs – 10 million years of evolution more different!
Considering their physical form and reproductive biology, the Asiatic salamanders are clearly some of the most primitive tailed amphibians. They have three traits which are considered particularly ancient: external fertilisation of their eggs; an angular bone in their lower jaw; and large numbers of genetic structures in their cells called microchromosomes. Lungs are present in adult Asiatic salamanders, except in the genus Onychodactylus (the “clawed salamanders” – the only salamanders to live without lungs outside of the Plethodontidae family). Hynobiids may either be terrestrial (land-dwelling) for much for their lives or predominantly aquatic (water-dwelling) and, like the Paradactylodon salamanders, tend to breed in mountain streams where oxygen is abundant. This causes them to have very small lungs (or no lungs at all) because large lungs could make it difficult to avoid becoming overly buoyant and swept away by strong currents.
Asiatic salamanders fall into two major categories: stream-type (like the Paradactylodon salamanders) or pond-type. Stream-type species live in streams or close to streams, and their young (or larvae) develop in running water, whilst pond-type species live in humid lowlands, and their larvae develop in still water. Either way, Asiatic salamanders are dependent upon moist conditions, and water bodies for breeding, and therefore have a very limited ability to disperse over long distances. Factors such as desertification from Mongolia to Western Asia about 50 million years ago, and uplift of the Tibetan plateau about 40 million years ago, and mountain generation (or orogeny) as the continents developed shaped the diversification and current distribution of the Asiatic salamanders today. All of these dramatic changes to the Asian landscape are thought to have been triggered by the collision of India and Asian about 50 million years ago. It is hypothesised that the first Asiatic salamanders originated in Northern China and were stream-adapted, and that their dispersal was subsequently affected by geography as they became restricted in their distribution by deserts, mountains and oceans.
The Paradactylodon salamanders speciated within the Asiatic salamanders about 40 million years ago in the Eocene period. They therefore arose 5 million years before the common ancestor of monkeys and humans. Found in the Central Asian countries of Iran and Afghanistan, they have the western-most distribution of all Asiatic salamanders, the rest being most heavily concentrated in eastern Asia. The Asiatic salamanders are an ancient lineage that has largely remained close to its likely site of origination in northern China. The three species Paradactylodon salamander in Central Asia are therefore minority outliers of this family in terms of their own distribution.
Considering their physical form and reproductive biology, the Asiatic salamanders are clearly some of the most primitive tailed amphibians. They have three traits which are considered particularly ancient: external fertilisation of their eggs; an angular bone in their lower jaw; and large numbers of genetic structures in their cells called microchromosomes. Lungs are present in adult Asiatic salamanders, except in the genus Onychodactylus (the “clawed salamanders” – the only salamanders to live without lungs outside of the Plethodontidae family). Hynobiids may either be terrestrial (land-dwelling) for much for their lives or predominantly aquatic (water-dwelling) and, like the Paradactylodon salamanders, tend to breed in mountain streams where oxygen is abundant. This causes them to have very small lungs (or no lungs at all) because large lungs could make it difficult to avoid becoming overly buoyant and swept away by strong currents.
Asiatic salamanders fall into two major categories: stream-type (like the Paradactylodon salamanders) or pond-type. Stream-type species live in streams or close to streams, and their young (or larvae) develop in running water, whilst pond-type species live in humid lowlands, and their larvae develop in still water. Either way, Asiatic salamanders are dependent upon moist conditions, and water bodies for breeding, and therefore have a very limited ability to disperse over long distances. Factors such as desertification from Mongolia to Western Asia about 50 million years ago, and uplift of the Tibetan plateau about 40 million years ago, and mountain generation (or orogeny) as the continents developed shaped the diversification and current distribution of the Asiatic salamanders today. All of these dramatic changes to the Asian landscape are thought to have been triggered by the collision of India and Asian about 50 million years ago. It is hypothesised that the first Asiatic salamanders originated in Northern China and were stream-adapted, and that their dispersal was subsequently affected by geography as they became restricted in their distribution by deserts, mountains and oceans.
The Paradactylodon salamanders speciated within the Asiatic salamanders about 40 million years ago in the Eocene period. They therefore arose 5 million years before the common ancestor of monkeys and humans. Found in the Central Asian countries of Iran and Afghanistan, they have the western-most distribution of all Asiatic salamanders, the rest being most heavily concentrated in eastern Asia. The Asiatic salamanders are an ancient lineage that has largely remained close to its likely site of origination in northern China. The three species Paradactylodon salamander in Central Asia are therefore minority outliers of this family in terms of their own distribution.
Asiatic salamanders are moderate to small in size, measuring less than 200 mm in length. The Paghman mountain salamander has 14 “costal” groves running across each side of its body. This species is known to possess two small arch-shaped rows of “vomerine” teeth, which are found on the roof of the mouth, and has a broadly rounded snout. The limbs are well-developed, and 4 fingers and toes are present with slightly overlapping digits. The tips of the digits are covered with a brown horny layer of skin that stops short of the soles and palms. A caudal fin is present along the tail, beginning just behind the hind legs with the highest point being about 2/3 of the way back. The tail slightly shorter than body. This species is dark olive-brown to yellowish olive in colouration, and the skin is indistinctly speckled with tiny pigmented dots. The belly is lighter, with fewer dots.
This species is completely aquatic (water-dwelling) and is probably active throughout the year because of the relatively constant flow of water in its inhabited streams. It only occurs within cold fast-flowing streams surrounded by arid scrub without trees, but in drier years these salamanders are restricted to the more constant environments of the three tributary sources of the Paghman stream drainage. Adults Paghman mountain salamanders are found under flat rocks, crawling through submerged scree rubble (or talus) or feeding along the stream bed. The young (or larvae) are usually found in deeper pools than the adults, out of the main flow of the current in areas shaded by vegetation, or under beds of submerged vegetation. During mating, the female produces paired arc-shaped, gelatinous egg sacs, each clutch containing 30-40 eggs. The male grasps these as they emerge from her cloaca (or reproductive opening) and, pressing them to his cloaca, sheds sperm onto them. The eggs of this species are then attached to the underside of rocks in the stream, although the total number of cases deposited by females is unknown.
Paghman mountain salamanders have a “biphasic life cycle” with aquatic larvae (similar to the tadpole stage in a frog’s development) that undergo metamorphosis into the adult form. The larvae have external gills, four pairs of gill slits, and a caudal (or tail) fin. These characteristics are largely lost at metamorphosis, and adult characteristics, such as eyelids and lungs, are acquired. Parental care of eggs is thought to occur through guarding by males, and the young stay in a larval state for 2-3 years.
Some Asiatic salamanders feed via tongue projection, rather like a chameleon (the genera Hynobius and Salamandrella). However, the rest of the species, including the Paradactylodon salamanders, take their food by “prehension” or sucking it from their watery environment. The larvae and adults prey mainly upon aquatic invertebrates, such as fresh-water shrimps, and the larvae of flies, caddis flies and beetles.
Paghman mountain salamanders have a “biphasic life cycle” with aquatic larvae (similar to the tadpole stage in a frog’s development) that undergo metamorphosis into the adult form. The larvae have external gills, four pairs of gill slits, and a caudal (or tail) fin. These characteristics are largely lost at metamorphosis, and adult characteristics, such as eyelids and lungs, are acquired. Parental care of eggs is thought to occur through guarding by males, and the young stay in a larval state for 2-3 years.
Some Asiatic salamanders feed via tongue projection, rather like a chameleon (the genera Hynobius and Salamandrella). However, the rest of the species, including the Paradactylodon salamanders, take their food by “prehension” or sucking it from their watery environment. The larvae and adults prey mainly upon aquatic invertebrates, such as fresh-water shrimps, and the larvae of flies, caddis flies and beetles.
This species is found in the Paghman Mountains of the Hindu Kush system, in the Paghman County of Afghanistan. The Paghman salamander inhabits cool highland streams fed by glaciers, where the adults are found to occur under rocks in fast-running water. The larvae live in deeper and quieter pools, commonly under floating beds of watercress and duckweed. Algae cover most of submerged rocks in the streams. The water temperature in summer varies between 0-14°C, and this species is found in waters above 14°C. This species is present between an altitudinal range of 2,750 and 3,050 metres above sea level, although it is increasingly uncommon in the lower elevations of this range. The vegetation surrounding the streams in which these salamanders occur is arid scrub without trees, and this species is known to be very sensitive to habitat disturbance.
This species is known from the three tributaries of the Paghman stream drainage system approximately 4km above the town of Paghman in the Hindu Kush of Kabul Province, Afghanistan. The stream is about 4 km in length and is fed by melting glaciers. The mid-point in the stream is located at 34º, 36' N, 68º 55' E. The salamanders are found in the altitudinal range of 2,440-3,750 metres above sea level.
Although the species was previously reported to be common between an altitudinal range of 2,750 and 3,050 metres above sea level, it now appears to be uncommon in the lower elevations of this range. The population is estimated to contain 1,000 to 2,000 adults.
This species is thought to be in decline by the IUCN Red List of Threatened Species, especially in the lower elevations of its range. The first formal collector, Mr. Chaworth-Musters, reported that the salamander was fairly common between the altitudes 2,750-3,050 metres above sea level, but observations made in 1977 and 1978 by S.M. Reilly revealed that the Paghman mountain salamander is no longer very common at these altitudes. It is now found towards the higher end of its known range. In general, the species seems to be highly endangered and unprotected, and its current status may only be speculated due to continuing civil war in Afghanistan.
The Paghman mountain salamander listed as Critically Endangered because its area of occupancy is less than 10 km sq., all individuals are in a single location, and there is continuing decline in the extent and quality of its habitat.
There is some evidence suggesting that there has been a constriction in range of this species at the lower elevations within its range, as well as a decline in the overall population. This alteration in the species’ distribution and numbers has probably been caused by irrigated cultivation close to its stream reducing flow. Additionally, grazing by livestock has removed stream edge vegetation causing an increase in temperature in the streams, since high water temperature may be an important factor limiting their local distribution as they are not found in water temperatures of above 14°C. Their habitat is also impacted by the physical disturbance of pedestrian and livestock (including sheep, goats, donkeys, horses and camels) traffic in and along the stream. A potential threat is the damming of the stream to provide a water source for the city of Kabul. The impact of the recent war in Afghanistan on this species is not known. The species has a very narrow distribution and ecological niche, which makes it highly vulnerable to habitat degradation, through a number of different threats, from climate change to interference with the flow of this species’ stream system.
The Paghman mountain salamander is not known from any protected areas, and there are currently no conservation measures underway fro this species, not least because of the ongoing civil unrest in Afghanistan.
An urgent priority for this species will be to conduct survey work as soon as the situation in Afghanistan permits. This will allow for an accurate assessment of this species’ population status to be conducted. Any information collected should be used to develop a Conservation Action Plan to galvanise future efforts for the preservation of this species in the wild. From what is known, it is clear that some habitat protection must occur for this species to prevent its untimely extinction. Conserving this stream system will require working with local people to encourage more ecologically sensitive land uses. The damming of this stream system to provide water for Kabul should also be addressed to maintain a minimum required level of stream flow for this species.
In addition to conserving native 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. Since the Paghman mountain salamander is categorised as Critically Endangered, the possibility of a captive breeding programme for this species should be investigated.
In addition to conserving native 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. Since the Paghman mountain salamander is categorised as Critically Endangered, the possibility of a captive breeding programme for this species should be investigated.
AmphibiaWeb: Information on amphibian biology and conservation [web application]. 2006. Berkeley, California: AmphibiaWeb. Available: amphibiaweb. Accessed: 08 December 2006.
Baillie, J. and Groombridge, B. (compilers and editors) 1996. 1996 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland.
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.
Groombridge, B. (ed.) 1994. 1994 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland.
Halliday, T. and Adler, C. (eds.). 2002. The new encyclopedia of reptiles and amphibians. Oxford University Press, Oxford.
Hilton-Taylor, C. (compiler). 2000. 2000 IUCN Red List of Threatened Species. IUCN, Gland, Switzerland and Cambridge, UK.
IUCN Conservation Monitoring Centre. 1988. 1988 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland and Cambridge, UK.
IUCN. 1990. 1990 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland and Cambridge, UK.
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 W. W. Dimmick. 1993. Phylogenetic relationships of the salamander families: A analysis of congruence among morphological and molecular characters. Herpetological Monographs 7:77-93.
Larson, A., D. W. Weisrock, and K. H. Kozak. 2003. Phylogenetic systematics of salamanders (Amphibia: Urodela), a review. Pp. 31-108 in Reproductive Biology and Phylogeny of Urodela (D. M. Sever, ed.) Science Publishers, Inc., Enfield (NH), USA.
Leviton, A.E. and Anderson, S.C. 1970. The amphibians and reptiles of Afghanistan a checklist and key to the herpetofauna. Proceedings of the California Academy of Sciences 38(10): 163-206.
Mertens, R. 1970. Salamander aus Afghanustan: Batrachuperus mustersi. DATZ 23: 346-348.
Nawabi, S. 1965. A rare amphibian from Afghanistan: Batrachuperus mustersi. Science (Kabul) (Aug. Sp. Iss.): 21-25.
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.
Papenfuss, T., Anderson, S. & Kuzmin, S. 2004. Batrachuperus mustersi. In: IUCN 2006. 2006 IUCN Red List of Threatened Species. IUCN Red List of Threatened Species.. Downloaded on 08 December 2006.
Reilly, S.M. 1983. The biology of the high altitude salamander Batrachuperus mustersi from Afghanistan. Journal of Herpetology 17(1): 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.
Smith, M.A. 1940. Contributions to the herpetology of Afghanistan. Ann. Mag. Nat. Hist. 11(5): 382-384.
Thorn, R. 1968. Les Salamandres d'Europe, d'Asia, et d'Afrique du Nord. Éditions Paul Lechevalier, Paris.
Zhang, P., Y.-Q. Chen, H. Zhou, X.-L. Wang, T. J. Papenfuss, D. B. Wake and L.-H. Qu. 2006. Phylogeny, evolution, and biogeography of Asiatic salamanders (Hynobiidae). Proceedings of the National Academy of Sciences 103:7360-7365.
Baillie, J. and Groombridge, B. (compilers and editors) 1996. 1996 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland.
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:
Groombridge, B. (ed.) 1994. 1994 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland.
Halliday, T. and Adler, C. (eds.). 2002. The new encyclopedia of reptiles and amphibians. Oxford University Press, Oxford.
Hilton-Taylor, C. (compiler). 2000. 2000 IUCN Red List of Threatened Species. IUCN, Gland, Switzerland and Cambridge, UK.
IUCN Conservation Monitoring Centre. 1988. 1988 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland and Cambridge, UK.
IUCN. 1990. 1990 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland and Cambridge, UK.
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 W. W. Dimmick. 1993. Phylogenetic relationships of the salamander families: A analysis of congruence among morphological and molecular characters. Herpetological Monographs 7:77-93.
Larson, A., D. W. Weisrock, and K. H. Kozak. 2003. Phylogenetic systematics of salamanders (Amphibia: Urodela), a review. Pp. 31-108 in Reproductive Biology and Phylogeny of Urodela (D. M. Sever, ed.) Science Publishers, Inc., Enfield (NH), USA.
Leviton, A.E. and Anderson, S.C. 1970. The amphibians and reptiles of Afghanistan a checklist and key to the herpetofauna. Proceedings of the California Academy of Sciences 38(10): 163-206.
Mertens, R. 1970. Salamander aus Afghanustan: Batrachuperus mustersi. DATZ 23: 346-348.
Nawabi, S. 1965. A rare amphibian from Afghanistan: Batrachuperus mustersi. Science (Kabul) (Aug. Sp. Iss.): 21-25.
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.
Papenfuss, T., Anderson, S. & Kuzmin, S. 2004. Batrachuperus mustersi. In: IUCN 2006. 2006 IUCN Red List of Threatened Species. IUCN Red List of Threatened Species.. Downloaded on 08 December 2006.
Reilly, S.M. 1983. The biology of the high altitude salamander Batrachuperus mustersi from Afghanistan. Journal of Herpetology 17(1): 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.
Smith, M.A. 1940. Contributions to the herpetology of Afghanistan. Ann. Mag. Nat. Hist. 11(5): 382-384.
Thorn, R. 1968. Les Salamandres d'Europe, d'Asia, et d'Afrique du Nord. Éditions Paul Lechevalier, Paris.
Zhang, P., Y.-Q. Chen, H. Zhou, X.-L. Wang, T. J. Papenfuss, D. B. Wake and L.-H. Qu. 2006. Phylogeny, evolution, and biogeography of Asiatic salamanders (Hynobiidae). Proceedings of the National Academy of Sciences 103:7360-7365.
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