Abstract
The phylogenetic relationships among members of the family Salamandridae have been repeatedly investigated over the last 90 years, with changing character and taxon sampling. We review the changing composition and the phylogenetic position of salamandrid genera and species groups and add a new phylogeny based exclusively on sequences of nuclear genes. Salamandrina often changed its position depending on the characters used. It was included several times in a clade together with the primitive newts (Echinotriton, Pleurodeles, Tylototriton) due to their seemingly ancestral morphology. The latter were often inferred as a monophyletic clade. Respective monophyly was almost consistently established in all molecular studies for true salamanders (Chioglossa, Lyciasalamandra, Mertensiella, Salamandra), modern Asian newts (Cynops, Laotriton, Pachytriton, Paramesotriton) and modern New World newts (Notophthalmus, Taricha). Reciprocal non-monophyly has been established through molecular studies for the European mountain newts (Calotriton, Euproctus) and the modern European newts (Ichthyosaura, Lissotriton, Neurergus, Ommatotriton, Triturus) since Calotriton was identified as the sister lineage of Triturus. In pre-molecular studies, their respective monophyly had almost always been assumed, mainly because a complex courtship behaviour shared by their respective members. Our nuclear tree is nearly identical to a mito-genomic tree, with all but one node being highly supported. The major difference concerns the position of Calotriton, which is no longer nested within the modern European newts. This has implications for the evolution of courtship behaviour of European newts. Within modern European newts, Ichthyosaura and Lissotriton changed their position compared to the mito-genomic tree. Previous molecular trees based on seemingly large nuclear data sets, but analysed together with mitochondrial data, did not reveal monophyly of modern European newts since taxon sampling and nuclear gene coverage was too poor to obtain conclusive results. We therefore conclude that mitochondrial and nuclear data should be analysed on their own.
Highlights
Salamandridae is the second speciose family of tailed amphibians
Freitag published what he called ‘his version’ of the phylogenetic relationships of the salamandrid genera based on morphology, life history and behaviour [15], whereas Laurent was the first who used dentition of jaws to discuss the phylogenetic relationships of Salamandridae [16]
Since we focus on the phylogenetic relationships of currently accepted genera, we did not extend our review to papers that studied intra-generic relationships, such as for the genus Salamandra (e.g., [22, 31, 32]) or the genus Triturus sensu lato (e.g., [33,34,35,36,37])
Summary
Salamandridae is the second speciose family of tailed amphibians. Currently, 21 genera with 118 species are recognized within three sub-families: Pleurodelinae (100 species in 16 genera; we here, in contrast to [1], consider Liangshantriton being part of Tylototriton, since [2], who introduced this new genus, did not provide a diagnosis [3], and the AmphibiaChina online database [4] does not list this genus), Salamandrinae (16 species in four genera) and Salamandrininae (2 species in one genus) ([1]; date of access: 10.10.2017). At that time, tailed amphibians (order Caudata) were subdivided into three suborders: Cryptobranchoidea, Ambystomoidea and Salamandroidea Within the latter, the family Salamandridae included today’s Salamandridae, and the genera Proteus, Necturus, Siren and Amphiuma [8]. Arnold only used courtship behaviour to infer his phylogeny of Salamandridae [13], while Naylor studied the vertebral column and the trunk musculature [14] Freitag published what he called ‘his version’ of the phylogenetic relationships of the salamandrid genera based on morphology, life history and behaviour [15], whereas Laurent was the first who used dentition of jaws to discuss the phylogenetic relationships of Salamandridae [16]. External morphology and mating behaviour, Scholz in 1995 inferred two alternative cladograms that differed only in the basal splits of true salamanders [17]
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