Roberto Sindaco

Conquering the Sahara and Arabian deserts: systematics and biogeography of Stenodactylus geckos (Reptilia: Gekkonidae)

BackgroundThe evolutionary history of the biota of North Africa and Arabia is inextricably tied to the complex geological and climatic evolution that gave rise to the prevalent deserts of these areas. Reptiles constitute an exemplary group in the study of the arid environments with numerous well-adapted members, while recent studies using reptiles as models have unveiled interesting biogeographical and diversification patterns. In this study, we include 207 specimens belonging to all 12 recognized species of the genus Stenodactylus. Molecular phylogenies inferred using two mitochondrial (12S rRNA and 16S rRNA) and two nuclear (c-mos and RAG-2) markers are employed to obtain a robust time-calibrated phylogeny, as the base to investigate the inter- and intraspecific relationships and to elucidate the biogeographical history of Stenodactylus, a genus with a large distribution range including the arid and hyper-arid areas of North Africa and Arabia.ResultsThe phylogenetic analyses of molecular data reveal the existence of three major clades within the genus Stenodactylus, which is supported by previous studies based on morphology. Estimated divergence times between clades and sub-clades are shown to correlate with major geological events of the region, the most important of which is the opening of the Red Sea, while climatic instability in the Miocene is hypothesized to have triggered diversification. High genetic variability is observed in some species, suggesting the existence of some undescribed species. The S. petrii - S. stenurus species complex is in need of a thorough taxonomic revision. New data is presented on the distribution of the sister species S. sthenodactylus and S. mauritanicus.ConclusionsThe phylogenetic hypothesis for the genus Stenodactylus presented in this work permits the reconstruction of the biogeographical history of these common desert dwellers and confirms the importance of the opening of the Red Sea and the climatic oscillations of the Miocene as major factors in the diversification of the biota of North Africa and Arabia. Moreover, this study traces the evolution of this widely distributed and highly specialized group, investigates the patterns of its high intraspecific diversity and elucidates its systematics.

Neogene climatic oscillations shape the biogeography and evolutionary history of the Eurasian blindsnake.

Typhlops vermicularis is the only extant scolecophidian representative occurring in Europe. Its main distribution area, the eastern Mediterranean, has a complicated geological and climatic history that has left an imprint on the phylogenies and biogeography of many taxa, especially amphibians and reptiles. Since reptiles are sensitive indicators of palaeogeographical and palaeoclimatic events, we investigated the intraspecific genealogy of T. vermicularis in a phylogeographical framework. A total of 130 specimens were analyzed, while the use of formalin and ethanol as preservatives called for a special treatment of the samples. Partial sequences of two mitochondrial (12S and ND2) and one nuclear (PRLR) marker were targeted and the results of the phylogenetic analyses (NJ, ML and BI) and the parsimony-network revealed the existence of 10 evolutionary significant units within this species. In combination with the results of the dispersal-vicariance analysis, we may conclude that the Eurasian blindsnake has encountered a sequence of extinction events, followed by secondary expansion from refugia. Estimation of divergence times showed that severe climatic changes between significantly wetter and drier conditions in the Late Neogene have played a key role on the evolutionary and biogeographical history of T. vermicularis. Additionally, both markers (mtDNA and nDNA) distinguished a largely-differentiated evolutionary lineage (Jordan and south Syria), which could even be reckoned as a full species. Our study reveals the existence of cryptic evolutionary lineages within T. vermicularis, which calls for further attention both on the protection of intraspecific varieties and the respective geographic areas that hold them.

Forgotten in the ocean: systematics, biogeography and evolution of the Trachylepis skinks of the Socotra Archipelago

Sindaco, R., Metallinou, M., Pupin, F., Fasola, M. & Carranza, S. (2012). Forgotten in the ocean: systematics, biogeography and evolution of the Trachylepis skinks of the Socotra Archipelago. —Zoologica Scripta, 41, 346–362.

Origin and in situ diversification in Hemidactylus geckos of the Socotra Archipelago

The Socotra Archipelago is an ancient continental fragment of Gondwanan origin and one of the most isolated landforms on Earth and a biodiversity hot spot. Yet, the biogeography and evolutionary history of its endemic fauna still remain largely overlooked. We investigate the origin, tempo and mode of diversification in the Hemidactylus geckos of the Socotra Archipelago. Concatenated and multilocus species coalescent analyses of Hemidactylus from Arabia and North Africa indicate that the Hemidactylus from Socotra do not form a monophyletic group and branch as three independent and well‐supported clades instead. Both the chronogram inferred using the gene tree approach of BEAST and the age‐calibrated multilocus species tree obtained using *BEAST suggest that the origin of Hemidactylus from Socotra may have involved a first vicariance event that occurred in the Early Miocene, followed by two independent transoceanic dispersal events that occurred more recently, during the Pliocene. Within Socotra, we analysed patterns of genetic diversity, the phylogeography and the demographic history in all seven nonintroduced species of Hemidactylus. Results based on two mitochondrial and two nuclear loci from 144 individuals revealed complex patterns of within‐island diversification and high levels of intra‐species genetic divergence. The interplay of both historical and ecological factors seems to have a role in the speciation process of this group of geckos. Interestingly, the case of H. forbesii and H. oxyrhinus, which inhabit the island of Abd al Kuri with an area of 133 km2, may represent one of the most extreme cases of intra‐island speciation in reptiles ever reported.

The global distribution of tetrapods reveals a need for targeted reptile conservation

The distributions of amphibians, birds and mammals have underpinned global and local conservation priorities, and have been fundamental to our understanding of the determinants of global biodiversity. In contrast, the global distributions of reptiles, representing a third of terrestrial vertebrate diversity, have been unavailable. This prevented the incorporation of reptiles into conservation planning and biased our understanding of the underlying processes governing global vertebrate biodiversity. Here, we present and analyse the global distribution of 10,064 reptile species (99% of extant terrestrial species). We show that richness patterns of the other three tetrapod classes are good spatial surrogates for species richness of all reptiles combined and of snakes, but characterize diversity patterns of lizards and turtles poorly. Hotspots of total and endemic lizard richness overlap very little with those of other taxa. Moreover, existing protected areas, sites of biodiversity significance and global conservation schemes represent birds and mammals better than reptiles. We show that additional conservation actions are needed to effectively protect reptiles, particularly lizards and turtles. Adding reptile knowledge to a global complementarity conservation priority scheme identifies many locations that consequently become important. Notably, investing resources in some of the world’s arid, grassland and savannah habitats might be necessary to represent all terrestrial vertebrates efficiently.The global distribution of nearly all extant reptile species reveals richness patterns that differ spatially from that of other taxa. Conservation prioritization should specifically consider reptile distributions, particularly lizards and turtles.

Out of Africa: Phylogeny and biogeography of the widespread genus Acanthodactylus (Reptilia: Lacertidae)

Acanthodactylus lizards are among the most diverse and widespread diurnal reptiles in the arid regions spanning from North Africa across to western India. Acanthodactylus constitutes the most species-rich genus in the family Lacertidae, with over 40 recognized species inhabiting a wide variety of dry habitats. The genus has seldom undergone taxonomic revisions, and although there are a number of described species and species-groups, their boundaries, as well as their interspecific relationships, remain largely unresolved. We constructed a multilocus phylogeny, combining data from two mitochondrial (12S, cytb) and three nuclear (MC1R, ACM4, c-mos) markers for 302 individuals belonging to 36 known species, providing the first large-scale time-calibrated molecular phylogeny of the genus. We evaluated phylogenetic relationships between and within species-groups, and assessed Acanthodactylus biogeography across its known range. Acanthodactylus cladogenesis is estimated to have originated in Africa due to vicariance and dispersal events from the Oligocene onwards. Radiation started with the separation into three clades: the Western and scutellatus clades largely distributed in North Africa, and the Eastern clade occurring mostly in south-west Asia. Most Acanthodactylus species diverged during the Miocene, possibly as a result of regional geological instability and climatic changes. We support most of the current taxonomic classifications and phylogenetic relationships, and provide genetic validity for most species. We reveal a new distinct blanfordii species-group, suggest new phylogenetic positions (A. hardyi, A. masirae), and synonymize several species and subspecies (A. lineomaculatus, A. boskianus khattensis and A. b. nigeriensis) with their phylogenetically closely-related species. We recommend a thorough systematic revision of taxa, such as A. guineensis, A. grandis, A. dumerilii, A. senegalensis and the pardalis and erythrurus species-groups, which exhibit high levels of intraspecific variability, and clear evidence of phylogenetic complexity.

Taxonomic reassessment of Blanus strauchi (Bedriaga, 1884) (Squamata: Amphisbaenia: Blanidae), with the description of a new species from south-east Anatolia (Turkey)

The study of mitochondrial and nuclear DNA sequences reveals that the polytypic Blanus strauchi is a species complex including three well-defined allopatric clades, one of which consists of two sub-clades. Only the two sub-clades of the Western clade are morphologically diagnosable in the field, whereas obvious characters to distinguish the Central and the Eastern clades are lacking. However, all four clades show significant statistical differentiation on meristic traits, as well as in morphometric characters of the head when compared by means of the geometric morphometrics. The genetic distance between the three major mitochondrial clades is comparable to the p-distances for the same markers observed between Blanus species-pairs from Morocco and the Iberian Peninsula, respectively. The nuclear marker confirms the mitochondrial clades, and shows that the three major clades do not share any haplotypes, as an indication of restricted gene flow among them. On the basis of this evidence, the taxonomy of Blanus strauchi is re-assessed: the Western clade corresponds to B. strauchi, with two subspecies: B. s. strauchi and B. s. bedriagae. The Central clade corresponds to B. aporus, here elevated at the species rank. For the eastern clade there are no available names, and therefore it is described here as Blanus alexandri sp. nov.

Unexpectedly High Levels of Cryptic Diversity Uncovered by a Complete DNA Barcoding of Reptiles of the Socotra Archipelago.

Few DNA barcoding studies of squamate reptiles have been conducted. Due to the significance of the Socotra Archipelago (a UNESCO Natural World Heritage site and a biodiversity hotspot) and the conservation interest of its reptile fauna (94% endemics), we performed the most comprehensive DNA barcoding study on an island group to date to test its applicability to specimen identification and species discovery. Reptiles constitute Socotra’s most important vertebrate fauna, yet their taxonomy remains under-studied. We successfully DNA-barcoded 380 individuals of all 31 presently recognized species. The specimen identification success rate is moderate to high, and almost all species presented local barcoding gaps. The unexpected high levels of intra-specific variability found within some species suggest cryptic diversity. Species richness may be under-estimated by 13.8–54.4%. This has implications in the species’ ranges and conservation status that should be considered for conservation planning. Other phylogenetic studies using mitochondrial and nuclear markers are congruent with our results. We conclude that, despite its reduced length (663 base pairs), cytochrome c oxidase 1, COI, is very useful for specimen identification and for detecting intra-specific diversity, and has a good phylogenetic signal. We recommend DNA barcoding to be applied to other biodiversity hotspots for quickly and cost-efficiently flagging species discovery, preferentially incorporated into an integrative taxonomic framework.

Phylogenetic position, origin and biogeography of Palearctic and Socotran blind-snakes (Serpentes: Typhlopidae)

The majority of the family Typhlopidae occurs in the Neotropic, Australasian, Indo-Malayan and Afrotropic ecoregions. They show a restricted distribution in the western Palearctic, where they include few native species, i.e. Rhinotyphlops simoni, R. episcopus and Typhlops vermicularis. A unique species among typhlopids is T. socotranus, found in Socotra, one of the most endemic-rich archipelagoes. In this study we determine the phylogenetic position of the above mentioned species and discuss their systematics, origin and biogeography. For this purpose we use three protein-coding nuclear markers (AMEL-amelogenin, BDNF-brain-derived neurotrophic factor and NT3-neurotrophin 3) to construct a time-calibrated phylogeny of the family Typhlopidae. Our results show that T. socotranus is a sister-species to T. vermicularis, while R. simoni and R. episcopus are sister-species to each other and are found within the African clade of the family, although they are geographically distributed in west Asia. Additionally we discuss several hypotheses on their origin, as well as the occurence of typhlopids in Eurasia.

Patterns of diversification in islands: A comparative study across three gecko genera in the Socotra Archipelago☆

In this study we used the complete fauna of geckos of the Socotra Archipelago to test whether the three gecko genera co-occurring in the islands (Pristurus, Hemidactylus and Haemodracon) produced similar outcomes of morphological and climatic diversification. To test this, we produced a time-calibrated tree of 346 geckos including all 16 endemic species of the archipelago and 26 potential close-relatives in the continent. Our dating estimates revealed that most of the diversity of geckos in the archipelago was the consequence of in situ diversification. However not all genera shared similar patterns of diversification. While in Hemidactylus and Haemodracon this involved great differences in body size and low levels of climatic diversification (mostly involving sympatric distributions), an opposite pattern appeared in Pristurus in which most of the diversification involved shifts in climatic envelopes (mostly involving allopatric and parapatric distributions) but almost no size differentiation. Consistently with this, Pristurus was the only genus in which rates of size diversification in islands were substantially lower than in the continent. This illustrates how different groups can greatly differ in their patterns of intra-island diversification and highlights the importance of taxon-dependent factors at determining different patterns of diversification in the same insular context.