Aim The intricate puzzle-like geography of the western Mediterranean is a product of long-term tectonic and orogenic events, supplemented by repeated climatic oscillations since the Pliocene. It offers numerous vicariance events that may be invoked to explain speciation in amphibians. We test for plausibility of two mutually exclusive Iberian-African vicariance hypotheses to explain the basal split within newts of the genus Pleurodeles: (1) the disconnection of the Betic arch c. 14 Ma and (2) the end of the Messinian salinity crisis 5.33 Ma.Location Specimens of P. waltl and P. poireti were sampled from 32 populations in Portugal, Spain, Morocco, Algeria and Tunisia.Methods Parts of three mitochondrial genes were sequenced (16S rRNA, cytochrome b, ATPase). Based on substitution rate constancy among lineages three different molecular clocks were calibrated to derive competing evolutionary scenarios for lineage evolution within Pleurodeles.Results One scenario was aligned with the dated fossil record and with historical events that are known to have enabled terrestrial faunal exchange between Europe and Africa. In Pleurodeles, such faunal exchange is more likely to have happened three times, resulting in the current pattern of species diversity and haplotype distribution: (1) following the disconnection of the Betic region from Iberia and connection of its southernmost part (present-day Rif Mountains) to Africa, c. 14 Ma; (2) closing of the Strait of Gibraltar prior to the Messinian salinity crisis, 5.59-5.33 Ma; (3) passive dispersal in recent times, caused by rafting on vegetation or inadvertent displacement by man. The results show that North African P. poireti populations comprise two distinct lineages; despite their geographical proximity, haplotype distribution within both lineages indicates totally different histories (range fragmentation vs. dispersal).Main conclusions Ribbed salamanders mainly evolved through allopatric speciation, driven by vicariance events. However, faunal exchange between Europe and Africa at the western end of the Mediterranean basin was linked to well-known events of physical contact between both continents. This sheds new light on the potential role of dispersal across marine barriers via rafting or even, presumably inadvertent, anthropogenic displacement for the initiation of speciation in amphibians.
The amphibian fauna of the Kingdom of Morocco was traditionally regarded as poor and closely related to its European counterpart. However, an increase in research during the last decades revealed a considerable degree of endemism amongst Moroccan amphibians, as well as phenotypic and genotypic inter- and intraspecific divergence. Despite this increase in knowledge, a comprehensible overview is lacking while several systematic issues have remained unresolved. We herein present a contemporary overview of the distribution, taxonomy and biogeography of Moroccan amphibians. Fourteen fieldtrips were made by the authors and colleagues between 2000 and 2012, which produced a total of 292 new distribution records. Furthermore, based on the results of the present work, we (i) review the systematics of the genus Salamandra in Morocco, including the description of a new subspecies from the Rif- and Middle Atlas Mountains, Salamandra algira splendens ssp. nov.; (ii) present data on intraspecific morphological variability of Pelobates varaldiiand Pleurodeles waltl in Morocco; (iii) attempt to resolve the phylogenetic position of Bufo brongersmai and erect a new genus for this species, Barbarophryne gen. nov.; (iv) summarize and assess the availability of tadpole-specific characteristics and bioacoustical data, and (v) summarize natural history data.
The keeping of exotic pets is currently under debate and governments of several countries are increasingly exploring the regulation, or even the banning, of exotic pet keeping. Major concerns are issues of public health and safety, animal welfare and biodiversity conservation. The keeping of reptiles and amphibians in captivity encompasses all the potential issues identified with keeping exotic pets, and many of those relating to traditional domestic pets. Within the context of risks posed by pets in general, the authors argue for the responsible and sustainable keeping of reptile and amphibian pets by private persons, based on scientific evidence and on the authors' own expertise (veterinary medicine, captive husbandry, conservation biology).
The Italian endemic genus Salamandrina was recently determined to be polytypic since two species, Salamandrina perspicillata and S. terdigitata, were recognized using both mitochondrial and nuclear genetic markers. However, data on their range is very scanty and no morphological traits are available to distinguish between these two taxa. A thorough field survey was carried out to detect and to sample new sites in areas in which Salamandrina remained unrecorded and to sample populations expected to occur in the documented range. Specimens were analyzed for variation of three mitochondrial DNA partial sequences of 12S and 16S ribosomal RNAs and cytochrome b genes. Attribution of individuals to either of the two species was therefore made on the basis of their haplotypes. Here we provide relevant information on the distribution of the two species, which were also found in areas where Salamandrina had previously been unrecorded, i.e. in the eastern Basilicata, in the region of Apulia and within the wide area which separated the two species. In the latter zone, a small area of sympatry between the two species was found. Furthermore, we analyzed the dorsal coloration pattern and size-related characters of the two species using Classification and Regression Tree analysis (CART), the results of which provided the first account of the morphological differences between Salamandrina perspicillata and S. terdigitata. The southern species, S. terdigitata, is smaller in size than S. perspicillata and differs in its coloration, mainly due to the red color on the dorsal surface of its tail.
The opening of the Gibraltar land bridge occurred at the end of the Messinian Salinity Crisis approximately 5.3 Mya, and was one of the main causes of vicariance between European and north-west African amphibians, resulting in the origin of several new species. However, little is currently known about the causes for post-Messinian amphibian differentiation in the Maghreb, although it is acknowledged that the Pleistocene glaciations probably had considerable influence on several species. The current study uses both species distribution modelling (MAXENT) and information from a total of 694 bp of mitochondrial data (351 from cytochrome b and 342 from 12S rRNA) from 36 representatives of all three recognized subspecies of Moroccan Salamandra to infer the phylogeny and biogeography of Salamandra algira tingitana, which is characterized by both viviparous and ovoviviparous populations. According to the results, the split between S. a. tingitana and S. a. algira from the Rif and Middle Atlas mountains took place approximately 1.6 Mya, and could have been caused by a shift towards a colder and drier climate that occurred during the upper Pliocene, which may have resulted in the isolation of Salamandra at increasingly higher altitudes, or in other climatically favourable areas. Several lineages within S. a. tingitana originated during the Pleistocene climatic oscillations, one of which gave rise to the viviparous populations north of the Oued Martil. It is suggested that the origin of viviparity in S. a. tingitana occurred during the last 600 000 years. In order to further understand the origin of the unique viviparous population of S. algira from North Africa, predictive distribution models of the viviparous and ovoviviparous populations of S. a. tingitana were created using MAXENT to assess environmental differences. Niche divergence was subsequently determined using Schoener's D and Warren et al.'s I niche similarity metrics. Predictive modelling and niche divergence analyses revealed significant environmental differences between the two reproductive types, which could have influenced the transition from ovoviviparity to viviparity.
The banded newt (genus Ommatotriton) is widely distributed in the Near East (Anatolia, Caucasus and the Levant) - an understudied region from the perspective of phylogeography. The genus is polytypic, but the number of species included and the phylogenetic relationships between them are not settled. We sequenced two mitochondrial and two nuclear DNA markers throughout the range of Ommatotriton. For mtDNA we constructed phylogenetic trees, estimated divergence times using fossil calibration, and investigated changes in effective population size with Bayesian skyline plots and mismatch analyses. For nuDNA we constructed phylogenetic trees and haplotype networks. Species trees were constructed for all markers and nuDNA only. Species distribution models were projected on current and Last Glacial Maximum climate layers. We confirm the presence of three Ommatotriton species: O. nesterovi, O. ophryticus and O. vittatus. These species are genetically distinct and their most recent common ancestor was dated at ∼25Ma (Oligocene). No evidence of recent gene flow between species was found. The species show deep intraspecific genetic divergence, represented by geographically structured clades, with crown nodes of species dated ∼8-13Ma (Miocene to Early Quaternary); evidence of long-term in situ evolution and survival in multiple glacial refugia. While a species tree based on nuDNA suggested a sister species relationship between O. vittatus and O. ophryticus, when mtDNA was included, phylogenetic relationships were unresolved, and we refrain from accepting a particular phylogenetic hypothesis at this stage. While species distribution models suggest reduced and fragmented ranges during the Last Glacial Maximum, we found no evidence for strong population bottlenecks. We discuss our results in the light of other phylogeographic studies from the Near East. Our study underlines the important role of the Near East in generating and sustaining biodiversity.
Lack of disease spill-over between adjacent populations has been associated with habitat fragmentation and the absence of population connectivity. We here present a case which describes the absence of the spill-over of the chytrid fungus Batrachochytrium salamandrivorans (Bsal) between two connected subpopulations of fire salamanders (Salamandra salamandra). Based on neutrally evolving microsatellite loci, both subpopulations were shown to form a single genetic cluster, suggesting a shared origin and/or recent gene flow. Alpine newts (Ichthyosaura alpestris) and fire salamanders were found in the landscape matrix between the two sites, which are also connected by a stream and separated by no obvious physical barriers. Performing a laboratory trial using alpine newts, we confirmed that Bsal is unable to disperse autonomously. Vector-mediated dispersal may have been impeded by a combination of sub-optimal connectivity, limited dispersal ability of infected hosts and a lack of suitable dispersers following the rapid, Bsal-driven collapse of susceptible hosts at the source site. Although the exact cause remains unclear, the aggregate evidence suggests that Bsal may be a poorer disperser than previously hypothesized. The lack of Bsal dispersal between neighbouring salamander populations opens perspectives for disease management and stresses the necessity of implementing biosecurity measures preventing human-mediated spread.
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