BackgroundThe pronounced morphological conservatism within Tarentola geckos contrasted with a high genetic variation in North Africa, has led to the hypothesis that this group could represent a cryptic species complex, a challenging system to study especially when trying to define distinct evolutionary entities and address biogeographic hypotheses. In the present work we have re-examined the phylogenetic and phylogeographic relationships between and within all Mediterranean species of Tarentola, placing the genealogies obtained into a temporal framework. In order to do this, we have investigated the sequence variation of two mitochondrial (12S rRNA and 16S rRNA), and four nuclear markers (ACM4, PDC, MC1R, and RAG2) for 384 individuals of all known Mediterranean Tarentola species, so that their evolutionary history could be assessed.ResultsOf all three generated genealogies (combined mtDNA, combined nDNA, and mtDNA+nDNA) we prefer the phylogenetic relationships obtained when all genetic markers are combined. A total of 133 individuals, and 2,901 bp of sequence length, were used in this analysis. The phylogeny obtained for Tarentola presents deep branches, with T. annularis, T. ephippiata and T. chazaliae occupying a basal position and splitting from the remaining species around 15.38 Mya. Tarentola boehmei is sister to all other Mediterranean species, from which it split around 11.38 Mya. There are also two other major groups: 1) the T. mauritanica complex present in North Africa and Europe; and 2) the clade formed by the T. fascicularis/deserti complex, T. neglecta and T. mindiae, occurring only in North Africa. The cladogenesis between these two groups occurred around 8.69 Mya, coincident with the late Miocene. Contrary to what was initially proposed, T. neglecta and T. mindiae are sister taxa to both T. fascicularis and T. deserti.ConclusionsAt least in the Iberian Peninsula and Northwest Africa, the lineages obtained have some geographic coherency, whilst the evolutionary history of the forms from Northeast Africa remains unclear, with a paraphyletic T. fascicularis with respect to T. deserti. The separation between the T. mauritanica complex and the clade formed by the T. fascicularis/deserti complex, T. neglecta and T. mindiae is coincident with the uplift of the Atlas Mountain chain, and the establishment of two distinct bioclimatic regions on each side of the barrier.

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The quantification of realized niche overlap and the integration of species distribution models (SDMs) with calibrated phylogenies to study niche evolution are becoming not only powerful tools to understand speciation events, but can also be used as proxies regarding the delimitation of cryptic species. We applied these techniques in order to unravel how the fundamental niche evolved during cladogenesis within the Tarentola mauritanica species-complex. Our results suggest that diversification within this complex, during the Miocene and Pleistocene, is associated with both niche divergence and niche conservatism, with a pattern that varies depending on whether the variables involved are related to the mean or seasonality of temperature and humidity. Moreover, climatic variables related to humidity and temperature seasonality were involved in the niche shift and genetic diversification of the European/North African clade during the Pleistocene and in its maintenance in a fundamental niche distinct from that of the remaining members of the group. This study further highlights the need for a taxonomic revision of the T. mauritanica species-complex.

:   Hemogregarines are the most-commonly reported hemoparasites in reptiles. In this work we analyzed samples from 572 individuals of 6 species of the wall gecko genus Tarentola from European and African countries adjacent to the Mediterranean Sea as well as from the Macaronesian islands. Screening was done using hemogregarine-specific primers for the 18S rRNA gene. Positive amplifications were sequenced so that the diversity of the hemogregarines from these hosts could be assessed within a phylogenetic framework. The results from the phylogenetic analysis showed that within Tarentola, the detected parasites are comprised of at least 4 distinct main lineages of Hepatozoon spp. In clades A and B, the new sequences clustered closely together with the ones previously known from individuals of the genus Tarentola and other species of geckos but also with those from other vertebrate host groups including skinks, snakes, iguanids, and rodents. Clade C included a sample from Tarentola angustimentalis of the Canary Islands. This sequence is the first molecular characterization of these hemogregarines in this archipelago. Until now, this lineage had only been found in lacertids, skinks, and snakes, so this infection extends the host range for this clade. Lastly, in the newly detected clade D, the retrieved parasite sequences form a group currently identified as exclusive of geckos. Our results show that geckos of Tarentola spp. harbor a great diversity of hemogregarines but also that further sampling and other tools, including a multi-locus approach using faster-evolving genetic markers, and identification of definitive hosts are needed to better understand the biology, diversity, and distribution of these parasites.