It takes two to tango – Phylogeography, taxonomy and hybridization in grass snakes and dice snakes (Serpentes: Natricidae: Natrix natrix, N. tessellata)

Using two mitochondrial DNA fragments and 13 microsatellite loci, we examined the phylogeographic structure and taxonomy of two codistributed snake species (Natrix natrix, N. tessellata) in their eastern distribution area, with a focus on Turkey. We found evidence for frequent interspecific hybridization, previously thought to be extremely rare, and for backcrosses. This underscores that closely related sympatric species should be studied together because otherwise the signal of hybridization will be missed. Furthermore, the phylogeographic patterns of the two species show many parallels, suggestive of a shared biogeographic history. In general, the phylogeographies follow the paradigm of southern richness to northern purity, but the dice snake has some additional lineages in the south and east in regions where grass snakes do not occur. For both species, the Balkan Peninsula and the Caucasus region served as glacial refugia, with several mitochondrial lineages occurring in close proximity. Our results show that the mitochondrial divergences in both species match nuclear genomic differentiation. Yet, in the former glacial refugia of grass snakes there are fewer nuclear clusters than mitochondrial lineages, suggesting that Holocene range expansions transformed the glacial hotspots in melting pots where only the mitochondrial lineages persisted, bearing witness of former diversity. On the other hand, the deep mitochondrial divergences in N. tessellata across its entire range indicate that more than one species could be involved, even though lacking microsatellite data outside of Turkey prevent firm conclusions. On the contrary, our microsatellite and mitochondrial data corroborate that N. megalocephala is invalid and not differentiated from sympatric populations of N. natrix. For Cypriot grass snakes, our analyses yielded conflicting results. A critical assessment of the available evidence suggests that N. natrix is a genetically impoverished recent invader on Cyprus and taxonomically not distinct from a subspecies also occurring in western Anatolia and the southern Balkans. Based on combined mitochondrial and nuclear genomic evidence we propose that for grass snakes the following subspecies should be recognized in our study region: (1) Natrix natrix vulgaris Laurenti, 1768, southeastern Central Europe and northern Balkans; (2) Natrix natrix moreoticus (Bedriaga, 1882), southern Balkans, western Anatolia, and Cyprus; and (3) Natrix natrix scutata (Pallas, 1771), eastern Anatolia, Caucasus region, Iran, northeastern distribution range (from eastern Poland and Finland to Kazakhstan and the Lake Baikal region). Thus, Natrix natrix cypriaca (Hecht, 1930) becomes a junior synonym of N. n. moreoticus and Natrix natrix persa (Pallas, 1814) becomes a junior synonym of N. n. scutata. Due to insufficient material, we could not resolve the status of Natrix natrix syriaca (Hecht, 1930) from the Gulf of İskenderun, southeastern Turkey.

Evolutionary history of the two North African hedgehogs (Mammalia: Erinaceidae) Atelerix algirus and Paraechinus aethiopicus based on phylogeography and species distribution modelling

Two species of hedgehogs are known to occur in northern part of Africa: the Algerian hedgehog Atelerix algirus and the Ethiopian hedgehog Paraechinus aethiopicus. Within each species several subspecies were described based on morphometrical data and pelage coloration, but all these subspecies have enigmatic and unclear definitions. We investigated the phylogeographical history and taxonomy of these two species based on mitochondrial DNA data covering the entire geographical distribution of A. algirus and the North African distribution of P. aethiopicus. We also used climatic niche modelling to make inferences about their evolutionary history. Low genetic diversity was recovered in both species. While no phylogeographic pattern was found in P. aethiopicus, two haplogroups were identified within A. algirus. This could be explained by the fact that continuous high or moderate climatic suitability occurred throughout most of the Saharan desert since the LGM (Last Glacial Maximum) for the first species, while during the LGM there were several disconnected areas of high climatic suitability for A. algirus: one in South-West Morocco, one at the coastal Moroccan-Algerian border and one in Tunisia-coastal Libya. Our genetic results confirm that A. algirus recently colonized Spain, Balearic and Canary Islands, and that this colonization was probably mediated by humans. Suitable climatic conditions occurred throughout most of the Southern and Eastern Iberian Peninsula during the last 6,000 years which could have favored the spatial expansion of the Algerian hedgehog after its arrival in Europe. According to our molecular results subspecific recognition within North Africa is unwarranted for both species.

Hidden tribe: A new species of Stream Toad of the genus Ansonia Stoliczka, 1870 (Anura: Bufonidae) from the poorly explored mountainous borderlands of western Thailand

The integrated results of morphological and molecular phylogenetic analyses confirmed the new species status of a recently discovered population of Ansonia from Suan Phueng District, Ratchaburi Province, Thailand. Ansonia karensp. nov. is separated from all other species of Ansonia by a unique combination of mensural, discrete morphological, and color pattern characteristics and is the sister species of A. thinthinae from Tanintharyi Division, Myanmar. This discovery fills a geographic hiatus of 350 km between it and A. kraensis from Ranong Province, Thailand. Ansonia karensp. nov. is the newest member of a long list of range-restricted endemics having been recently discovered in the northern Tenasserim Mountain region of western Thailand and continues to underscore the unexplored nature of this region and its need for conservation.

Hidden diversity within a polytypic species: The enigmatic Sceloporus torquatus Wiegmann, 1828 (Reptilia, Squamata, Phrynosomatidae)

The spiny lizard genus Sceloporus was described by Wiegmann in 1828, with S. torquatus posteriorly designated as the type species. The taxonomic history of S. torquatus is complicated, as it has been confused with other taxa by numerous authors. Many modern systematics works have been published on Sceloporus, but none have included all five recognized S. torquatus subspecies: S. t. torquatus, S. t. melanogaster, S. t. binocularis, S. t. mikeprestoni, and S. t. madrensis. Additionally, there is previous evidence for at least one unnamed taxon. The present study is the first taxonomic revision of the enigmatic S. torquatus based on molecular phylogenies using combined molecular data from 12S, ND4 and RAG1 genes, and Maximum Likelihood and Bayesian inference phylogenetic methods. This work includes the most extensive sampling across the entire distribution, as well as divergence time estimates and environmental niche modelling, which combined offer a spatio-temporal framework for understanding the evolution of the species. Additionally, a series of morphological characters are analyzed to identify significant differences between lineages consistently recovered in the molecular phylogenies. Using this integrative approach, evidence is presented for eight lineages within the S. torquatus complex, five of which correspond to previously recognized subspecies and three represent unnamed taxa masked by morphological conservatism. Finally, to maintain taxonomic stability a lectotype and paralectoype are designated for S. torquatus, and certain taxonomic changes are suggested in order to reflect the phylogenetic relationships within the S. torquatus complex.

Phylogenetic relationships of xenodermid snakes (Squamata: Serpentes: Xenodermidae), with the description of a new genus

Xenodermidae is a generally poorly known lineage of caenophidian snakes found in South, East and Southeast Asia. We report molecular phylogenetic analyses for a multilocus data set comprising all five currently recognised genera and including new mitochondrial and nuclear gene sequence data for the recently described Stoliczkia vanhnuailianai. Our phylogenetic results provide very strong support for the non-monophyly of Stoliczkia, as presently constituted, with S. borneensis being more closely related to Xenodermus than to the Northeast Indian S. vanhnuailianai. Based on phylogenetic relationships and morphological distinctiveness, we transfer Stoliczkia borneensis to a new monotypic genus endemic to Borneo, Paraxenodermusgen. nov. We also present new morphological data for P. borneensis.

A new cryptic arboreal species of the Cyrtodactylus brevipalmatus group (Squamata: Gekkonidae) from the uplands of western Thailand

The first integrative taxonomic analysis of the Cyrtodactylus brevipalmatus group of Southeast Asia recovered two newly discovered populations from the Tenasserim Mountains in Suan Phueng District, Ratchaburi Province, Thailand as a new species described here as C. rukhadeva sp. nov. Based on 1397 base pairs of the mitochondrial gene NADH dehydrogenase subunit 2 (ND2), C. rukhadeva sp. nov. is the well-supported sister species to a clade containing three undescribed species, C. ngati, and C. cf. interdigitalis with a large uncorrected pairwise sequence divergence from other species in the brevipalmatus group ranging from 15.4–22.1%. Cyrtodactylus elok and C. brevipalmatus are recovered as poorly supported sister species and the well-supported sister lineage to the remainder of the brevipalmatus group. Cyrtodactylus rukhadeva sp. nov. is putatively diagnosable on the basis of a number of meristic characters and easily separated from the remaining species of the brevipalmatus group by a number of discrete morphological characters as well as its statistically significant wide separation in multivariate morphospace. The discovery of C. rukhadeva sp. nov. continues to underscore the unrealized herpetological diversity in the upland forests of the Tenasserim Mountains and that additional field work will undoubtedly result in the discovery of additional new species.

A new species of Myotis (Chiroptera, Vespertilionidae) from Uruguay

The genus Myotis comprises a diverse group of vesper bats with worldwide distribution. Twenty-eight neotropical species are currently recognized. Based on a morphological approach, we describe a new species of Myotis from the Uruguayan Pampas grasslands, an ecoregion under high anthropogenic pressure with a largely unknown bat fauna. Qualitative and quantitative morphological analyses support the recognition of the new species and we present a set of external and cranial diagnostic characters by comparing them with other neotropical Myotis species. The new species reassembles Myotis riparius, but can be distinguished by a set of qualitative and quantitative morphological traits, including its clearly bicolored dorsal fur, tricolored ventral fur, a pelage on the dorsal surface of uropatagium, sagittal crest lower, braincase lower in lateral view and overall smaller size.

Molecular phylogenetics and taxonomic reassessment of the widespread agamid lizard Calotes versicolor (Daudin, 1802) (Squamata, Agamidae) across South Asia

The genus Calotes Cuvier, 1817 (Agamidae: Draconinae) is highly diverse, with species occurring in South and Southeast Asia, and Oceania. Most species of the subfamily except C. versicolor have narrow geographic distributions. Calotes versicolor is distributed from western Iran in the west to south China and Indonesia in the east and has been introduced to parts of Africa and North America. The species has had a complicated taxonomic history; multiple species and subspecies related to C. versicolor were described from India and adjoining regions, which were synonymized in subsequent revisions. However, a study of Burmese C. versicolor yielded two new species, C. htunwini and C. irawadi, indicating that C. versicolor is a species complex. Such integrative taxonomic studies have not been carried out in India, the supposed type locality of C. versicolor. Hence, we studied C. versicolor sensu lato from the Indian subcontinent and generated sequences of mitochondrial 16S and COI fragments from tissues sampled from multiple localities in the region, including the type localities of its synonyms. Phylogenetic analyses revealed four well-supported, deeply-divergent lineages, supported by morphological data. These lineages represent (i) C. versicolor sensu stricto, from South India and parts of the east coast, (ii) C. irawadi sensu lato from northeast India and Southeast Asia, (iii) a synonym from the eastern Indo-Gangetic Plains which we resurrect here, and (iv) a subspecies from Pakistan which we elevate to species level. We provide re-descriptions for the resurrected or elevated species, and a diagnostic key to the species of the C. versicolor complex. The study shows that C. versicolor sensu stricto is endemic to parts of southern and eastern India, and not widely distributed, though it may have been introduced to other parts of the world.

Osteology of Tucanoichthys tucano Géry and Römer, an enigmatic miniature fish from the Amazon basin, Brazil (Teleostei: Characiformes: Characidae)

Miniaturization, the evolution of extremely small adult body size, is a common phenomenon across the lineages of freshwater fishes, especially in the Neotropics where over 200 species are considered miniature (≤26 mm in standard length [SL]). Close to 30% of all miniature Neotropical freshwater fishes belong to the family Characidae, several of which are of uncertain phylogenetic placement within the family. We investigate the skeletal anatomy of Tucanoichthys tucano, a species of uncertain phylogenetic position from the upper Rio Negro basin, reaching a maximum known size of 16.6 mm SL. The skeleton of Tucanoichthys is characterized by the complete absence of ten skeletal elements and marked reduction in size and/or complexity of others, especially those elements associated with the cephalic latero-sensory canal system. Missing elements in the skeleton of Tucanoichthys include those that develop relatively late in the ossification sequence of the non-miniature characiform Salminus brasiliensis, suggesting that their absence in Tucanoichthys can be explained by a simple scenario of developmental truncation. A number of the reductions in the skeleton of Tucanoichthys are shared with other miniature characiforms, most notably species of Priocharax and Tyttobrycon, the latter a putative close relative of Tucanoichthys based on molecular data.

Humans, megafauna and landscape structure – Rock engravings from Namibia encourage a comparative approach to central Europe and southern Africa

This paper deals with reflections that arose after observing prehistoric rock engravings at different locations in Namibia. These observations stimulated comparative considerations with focus on southern Africa and central Europe. Similar to the Aurignacian rock art of European origin, the most common motifs in the Namibian rock engravings are large animals. While in Europe, the species that served as a blueprint for the illustration of Aurignacian rock art have mostly disappeared, the megafauna illustrated on the rock engravings in Namibia can still be found in the immediate vicinity of the rock art. Against this background, we discuss and further develop a comparative regional approach. We reconstruct and evaluate the suitability of African savannas and still-existing megafauna communities as an appropriate reference-frame for natural European grassland systems and extinct associated warm-adapted megafauna (Eemian Interglacial megafauna). Special attention is laid on the unique situation in Africa in the light of a global extinction wave of megafauna following increasing human activity in the Late Quaternary. This leads us to discuss the use of domesticated ungulates as surrogate taxa to fulfill ecosystem functions in Europe as part of concepts termed “rewilding” or “naturalistic grazing”. After critically examining these concepts, we conclude that using domesticated forms as representatives of extinct or locally disappeared species in Europe has its justification to some extent. If, however, the naturally occurring megaherbivore community still exists (Africa), these naturally occurring species should be given priority due to their organismic abilities and limitations adapted to the harsh conditions in their specific environment. Finally, we discuss the application of (transboundary) protected areas as effective instruments to mitigate human-wildlife conflicts. A holistic approach, including nature conservation and preservation of cultural achievements (domesticated forms, grazing systems), appears promising for the effective protection of the natural African savanna ecosystems with their unique fauna elements, as illustrated in rock engravings that inspired us to write this paper.