47.
Gorgan Mountain Salamander
(Paradactylodon gorganensis)
CR
Overview
The Gorgan mountain salamander is restricted to a single cave and stream in the mountains of northern Iran. It lives almost entirely in water, feeding on invertebrates and breeding in a long, narrow pool within the Shir-Abad cave. During mating, the female produces paired arc-shaped, gelatinous egg sacs, each containing 35-70 eggs. The male grasps these and fertilises them externally. There are only around 100 breeding adults remaining, and all are threatened by a restricted distribution and numerous visitors each year to the Shir-Abad cave. The is some question regarding whether the Gorgan mountain salamander is a separate species, or a local population of the Persia mountain salamander (Paradactylodon persicus).
Urgent Conservation Actions
Prevent tourists from damaging the Shir-Abad cave; resolve the species status of the Gorgan mountain salamander.
Distribution
Golestan Province, northern Iran
Fact
The Gorgan mountain salamander may not be a separate species, but rather a local population of the Persia mountain salamander (Paradactylodon persicus).
Evolutionary Distinctiveness
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.
Description
Asiatic salamanders are moderate to small in size, measuring less than 200 mm in length. The Gorgan mountain salamander was originally described from just a single male individual in 1979. This species is known to possess two small arch-shaped rows of “vomerine” teeth, which are found on the roof of the mouth. It has a large and flattened head with a rounded snout. The limbs, especially the hind limbs, muscular and robust. Four toes and fingers are present and they do hot have “claws” or keratinised digit tips like some species of Asiatic salamander. The tail is longer than the body, and is flattened from the sides and tapers into a rounded point. The skin is smooth, with numerous pores and is yellowish in colour, although this may not truly reflect the colour in life because this species was described from a specimen that had been preserved in alcohol. The unmetamorphosed young (or larvae) have external gills, four pairs of gill slits and caudal (or tail) fins.
Ecology
The Gorgan mountain salamander is an almost fully aquatic (water-dwelling) species. The adults are only known from a single pool in the cave in Iran’s Shir-Abad cave that measures 100m by 10m at its widest point. The larvae of this species may be found outside of the cave in the stream flowing from this lake. The breeding biology of this species is not well established, but eggs are known to be deposited by the female and fertilised externally by the male. During mating, the female produces paired arc-shaped, gelatinous egg sacs, each containing 35-70 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 sacs may then be attached to stones or vegetation in the water.

Gorgan 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 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.

The feeding ecology of the Gorgan mountain salamander is almost completely unknown. Insect remains were found in the digestive tract of the individual that was originally described. 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.
Habitat
The Gorgan mountains salamander was discovered in the eastern part of the Elburz Mountains, in Iran’s Shir-Abad cave. The cave is located on the wetter, northern slope of the ridge, which is exposed to the Caspian Sea. Adults of this species have only been found in the Shir-Abad cave, which has a length of 200 m and contains a 100 m long pool with a stream exiting at the end. The larvae of this species may be found outside of the cave in this stream. The forest surrounding the cave and stream is a temperate rainforest type called Hyrcanian forest.

Adult Gorgan mountain salamander have been found inside this cave at the border of the stream. It is supposed that this species is trogloxene, i.e. a species not completely adapted to life in caves but one that lives closely associated with cave habitats nonetheless. The larvae of this species may be found outside of the cave in the stream.
Distribution
This species is only found in the Shir-Abad Cave and the stream flowing from it, 60 km east of Gorgan (36 57' N, 55 01' E), in the eastern part of the Elburz Mountains, in the Golestan Province, northern Iran. The cave is at an elevation of 310 metres above sea level.
Population Estimate
There are estimated to be about 100 breeding adults within its extremely localised range. However, the Gorgan mountain salamander is thought by some researchers not to be a separate species in its own right, but rather a local population of the Persia mountain salamander (Paradactylodon persicus).
Population Trend
This species is thought to be in decline by the IUCN Red List of Threatened Species.
Status
The Gorgan mountain 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. and 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 in the Shir-Abad Cave, northern Iran.
Threats
The Shir-Abad Cave is now significantly impacted by the activities of people visiting the cave. However, the forest surrounding both the cave and stream does not appear to have been logged, since the area is generally considered to be too rugged and remote for development. The Gorgan mountain salamander is highly specialised in terms of its habitat and has a very narrow range. Although trends and threats are largely unknown, it is evident that this species is vulnerable to habitat degradation as a result of its highly limited distribution.
Conservation Underway
Almost the whole of the Hyrcanian forests of Iran are regarded as protected areas, including the range of this species. Shir-Abad Cave and the surrounding area was designated a Natural National Place by the Department of Environment of Gorgan and Gonbad-e-Kavous in 1998.
Conservation Proposed
It is important to conserve the only known habitat of this species, and to minimise the damage done to Shir-Abad cave by visitors. Continued effective conservation of the area is also a major priority to insure that the only water source of the Gorgan mountain salamanders is not polluted or reduced. It may also be an important consideration to study this species in more detail to determine whether it is indeed a distinct species, or whether it is a localised population or subspecies if the Persia mountain salamander (Paradactylodon persicus).

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 Gorgan mountain salamander is categorised as Critically Endangered, the possibility of a captive breeding programme for this species should be investigated.
Links
References
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Clergue-Gazeau, M. and Thorn, R. 1979. Une nouvelle espece de salamandre du genre Batrachuperus en provenence de l'Iran septentrional (Amphibia, Caudata, Hynobiidae). Bulletin de la Société d'Histoire Naturelle de Toulouse 114: 455-460.

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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.

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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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