Introduction

Biology of salamanders Amphibians are a class of ‘cold­ adult life in water (e.g. the genera Cynops, Euproctus, Lissotriton, blooded’ vertebrate animals that have a naked, glandular skin, Neurergus, Triturus, and the American Taricha and Notophthalmus). which is lacking any epidermal covering such as fur, scales or There is however no defi nitive division between species called feathers. Salamanders are tailed amphibians with an elongated either newts or salamanders and the literature is not consistent body, prominent tail, and typically two pairs of limbs that are in its usage of the terms, in particular with regard to the Asian similar in size. The front legs have four fi ngers and the hind legs species. In other European languages the situation is similar. have fi ve or four. Salamanders belong to the order Caudata Throughout this book the term ‘salamander’ will be used to (‘caudate’ meaning ‘tailed’), which is one of the three orders refer to all species in the family Salamandridae, except for those comprising the class Amphibia. The other amphibian orders that have a partly aquatic lifestyle and which are commonly are the Anura (frogs and toads which lack tails and have bodies called ‘newt’ in English, ‘Molch’ in German or ‘triton’ in French, and limbs that are modifi ed for jumping) and the Gymnophiona which I will refer to as ‘newts’. (caecilians, a group of legless, worm­like animals found in the tropics). Because of their long bodies and tails, salamanders Currently 652 salamander species are recognised according superfi cially resemble lizards, which are reptiles. But unlike to Amphibian Species of the World (http://research.amnh.org/vz/ lizards, salamanders do not have scales or claws and have herpetology/amphibia/ accessed on 15 May 2013). They occur in aquatic larvae. Salamanders and lizards are very different Europe, in North, Central and South America, in Northern animals with different life histories and live in different Africa, and in Asia. Salamanders vary in length between 3 cm habitats. and 150 cm. By far the majority of species (433) belong to the family of lungless salamanders (Plethodontidae). These are Strictly speaking, all tailed amphibians are ‘salamanders’, found mainly in North, Central and South America, but eight and ‘newts’ are just a subset of the group. Hence, newts are species occur in Europe: the European cave salamanders of the salamanders, but not every salamander is a newt. ‘Newt’ is the genera Speleomantes and Atylodes. Much to the surprise of the common term generally used for those salamander species in scientifi c world, a new plethodontid salamander was discovered the family Salamandridae that spend at least part of their in South Korea in 2005 and named Karsenia koreana.

Caudata Cryptobranchidae Andrias A B Cryptobranchus Cryptobranchidae Hynobiidae Onychodactylus Sirenidae Pachyhynobius Salamandrella Salamandridae Hynobius Ambystomatidae Paradactylodon Hynobiidae Ranodon Dicamptodontidae Batrachuperus Proteidae Liua Pseudohynobius Rhyacotritonidae Salamandrina Amphiumidae Chioglossa Mertensiella Plethodontidae Lyciasalamandra Salamandra Pleurodeles Echinotriton Tylototriton Notophthalmus Taricha Lissotriton Salamandridae Calotriton Triturus Neurergus Phylogenetic relationships of extant salamanders based on Ommatotriton Euproctus molecular data. The branches of the tree represent lines of Ichthyosaura descent of salamander families of the world (A) and of genera in Laotriton Pachytriton the families Cryptobranchidae, Hynobiidae and Salamandridae Cynops (B). Tree topology follows Pyron & Wiens (2011). Paramesotriton

14 Salamanders of the Old World | Introduction The family Salamandridae is a group of salamanders that occur in America and are not part of the Old World salamander occurs mainly in Europe and Asia comprising just over fauna. 100 species, as well as two American newt genera, Taricha and Notophthalmus. The typical European and Asian newts The family of hynobiid salamanders (Hynobiidae) contains (Lissotriton and Cynops) and the Fire Salamander (Salamandra 59 species and only occurs in Asia, with the exception of the salamandra) belong to this family. Two other salamander Siberian Salamander (Salamandrella keyserlingii) which just families, the mole salamanders, Ambystomatidae (37 species reaches into Europe. In contrast to the other families, fertili­ of which the Axolotl, Ambystoma mexicanum, is the best known sation in the Hynobiidae and Cryptobranchidae is external, representative) and the small family Rhyacotritonidae only outside the body of the female. In hynobiids the eggs are

the only westerner who was allowed by the Japanese government to collect cultural and biological objects in Japan at that time. Von Siebold was employed as a medical doctor at the VOC (United East­Indian Company) trading post Deshima. In the early 19th century the activities of Von Siebold and his assistant Heinrich Bürger provided the only reliable source of knowledge about Japanese nature and culture. Von Siebold’s notes, the living specimen and a skeleton were used by biologist Coenraad Jacob Temminck to describe the Giant Salamander as a new species. Von Siebold’s notes and collection were the basis of two com­ prehensive publications about the natural history of Japan. Researchers of the State Museum of Natural History in Leiden published the book series Fauna Japonica, while researchers at the State Herbarium in Leiden wrote the Flora Japonica. Fauna ‘Salamandra maxima’, the Japanese Giant Salamander Japonica was published in six volumes between 1833 and 1850 and © Naturalis Biodiversity Center. Photo: Max Sparreboom. was produced by C. J. Temminck (1778­1858), Hermann Schlegel (1804­1884) and Willem de Haan (1801­1855). The Japanese Giant Salamander was described in Volume 3 and depicted with the The most famous illustration of the Japanese Giant Salamander beautiful lithograph shown here. (now called Andrias japonicus) is the lithograph published as plate VII of the Amphibians and Reptiles volume of the Fauna Japonica. The salamander shown in the picture was actually one of two The original is a coloured drawing made by Dr S. Mulder from specimens shipped by Von Siebold. The other ended up in the a living specimen and is kept in the library of the Naturalis stomach of the fi rst during the long sea voyage! Upon its arrival Biodiversity Center in Leiden. The plate was in a poor state of in Amsterdam, the salamander was fi rst taken to the State preservation and was restored in 2004. This photo was made Museum of Natural History in Leiden, where it was kept alive after the restoration. for 10 years. In 1840 it was given to the Amsterdam Zoo, Artis, On the original drawing, the salamander is depicted life­size. where it eventually died in 1881. The dead animal was preserved By positioning the salamander with head and tail bent in an in alcohol in a bottle, but the bottle appears to be lost. s­shape, the artist could fi t it onto a single page, while maintain­ Fortunately Von Siebold sent other specimens to the Nether­ ing enough detail of the skin structure. The illustration is used lands. These have been registered as paralectotypes in the in many later works on the Giant Salamander and even in logos of scientifi c collections of the Naturalis Biodiversity Center in Giant Salamander research and education institutions in Japan. Leiden and the zoological museums in Berlin and Paris. This drawing depicts the fi rst living Giant Salamander ever seen outside Japan. The animal was shipped from Nagasaki to Amster­ References dam in 1830 along with other animals, plants and Japanese Gassó Miracle et al. (2007); Hoogmoed (1978); Holthuis & Sakai (1970); utensils. It was collected by Philipp Franz von Siebold (1796­1866), Schlegel (1842); Temminck (1836); Temminck & Schlegel (1838).

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