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The changing views on the evolutionary relationships of extant Salamandridae (Amphibia: Urodela)
(2018)
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.
This thesis is focused on improving the knowledge on a group of threatened species, the European cave salamanders (genus Hydromantes). There are three main sections gathering studies dealing with different topics: Ecology (first part), Life traits (second part) and Monitoring methodologies (third part). First part starts with the study of the response of Hydromantes to the variation of climatic conditions, analysing 15 different localities throughout a full year (CHAPTER I; published in PEERJ in August 2015). After that, the focus moves on identify which is the operative temperature that these salamander experience, including how their body respond to variation of environmental temperature. This study was conducted using one of the most advanced tool, an infrared thermocamera, which gave the opportunity to perform detailed observation on salamanders body (CHAPTER II; published in JOURNAL OF THERMAL BIOLOGY in June 2016). In the next chapter we use the previous results to analyse the ecological niche of all eight Hydromantes species. The study mostly underlines the mismatch between macro- and microscale analysis of ecological niche, showing a weak conservatism of ecological niches within the evolution of species (CHAPTER III; unpublished manuscript). We then focus only on hybrids, which occur within the natural distribution of mainland species. Here, we analyse if the ecological niche of hybrids shows divergences from those of parental species, thus evaluating the power of hybrids adaptation (CHAPTER IV; unpublished manuscript). Considering that hybrids may represent a potential threat for parental species (in terms of genetic erosion and competition), we produced the first ecological study on an allochthonous mixed population of Hydromantes, analysing population structure, ecological requirements and diet. The interest on this particular population mostly comes by the fact that its members are coming from all three mainland Hydromantes species, and thus it may represent a potential source of new hybrids (CHAPTER V; accepted in AMPHIBIA-REPTILIA in October 2017). The focus than moves on how bioclimatic parameters affect species within their distributional range. Using as model species the microendemic H. flavus, we analyse the relationship between environmental suitability and local abundance of the species, also focusing on all intermediate dynamics which provide useful information on spatial variation of individual fitness (CHAPTER VI; submitted to SCIENTIFIC REPORTS in November 2017). The first part ends with an analysis of the interaction between Hydromantes and Batracobdella algira leeches, the only known ectoparasite for European cave salamanders. Considering that the effect of leeches on their hosts is potentially detrimental, we investigated if these ectoparasites may represent a further threat for Hydromantes (CHAPTER VII; submitted to INTERNATIONAL JOURNAL FOR PARASITOLOGY: PARASITES AND WILDLIFE in November 2017). The second part is related to the reproduction of Hydromantes. In the first study we perform analyses on the breeding behaviour of several females belonging to a single population, identifying differences and similarities occurring in cohorting females (CHAPTER VIII; published in NORTH-WESTERN JOURNAL OF ZOOLOGY in December 2015). In the second study we gather information from all Hydromantes species, analysing size and development of breeding females, and identifying a relationship between breeding time and climatic conditions (CHAPTER IX; submitted to SALAMANDRA in June 2017). In the last part of this thesis, we analyse two potential methods for monitoring Hydromantes populations. In the first study we evaluate the efficiency of the marking method involving Alpha tags (CHAPTER X; published in SALAMANDRA in October 2017). In the second study we focus on evaluating N-mixtures models as a methodology for estimating abundance in wild populations (CHAPTER XI; submitted to BIODIVERSITY & CONSERVATION in October 2017).