Wolfgang Böhme

Phylogenetic Affinities of Mabuya atlantica Schmidt, 1945, Endemic to the Atlantic Ocean Archipelago of Fernando de Noronha (Brazil): Necessity of Partitioning the Genus Mabuya Fitzinger, 1826 (Scincidae: Lygosominae)

Abstract The infrageneric phylogenetic position of the Brazilian skink Mabuya atlantica was inferred from 859 bp of the mitochondrial 16S and 12S rRNA genes. It could be shown that M. atlantica belongs to the Afro-Malagasy rather than to the South American Mabuya radiation. Mabuya atlantica probably represents an independent transmarine colonization directly from the coast of Southwest Africa, thus representing another example of the extraordinary dispersal abilities of members of this group. Moreover, the present analysis revealed that intercontinental relationships within the genus Mabuya are far more complex than previously thought. The molecular analysis suggests that Mabuya consists of several long-separated evolutionary lineages, representing distinct and well-supported monophyletic radiations. To reflect the independent origins of the South American, Asian, Afro-Malagasy and Cape Verdian groups we partition the genus Mabuya into four genera.

Eastward from Africa: palaeocurrent-mediated chameleon dispersal to the Seychelles islands

Madagascar and the Seychelles are Gondwanan remnants currently isolated in the Indian Ocean. In the Late Cretaceous, these islands were joined with India to form the Indigascar landmass, which itself then split into its three component parts around the start of the Tertiary. This history is reflected in the biota of the Seychelles, which appears to contain examples of both vicariance- and dispersal-mediated divergence from Malagasy or Indian sister taxa. One lineage for which this has been assumed but never thoroughly tested is the Seychellean tiger chameleon, a species assigned to the otherwise Madagascar-endemic genus Calumma. We present a multi-locus phylogenetic study of chameleons, and find that the Seychellean species is actually the sister taxon of a southern African clade and requires accomodation in its own genus as Archaius tigris. Divergence dating and biogeographic analyses indicate an origin by transoceanic dispersal from Africa to the Seychelles in the Eocene–Oligocene, providing, to our knowledge, the first such well-documented example and supporting novel palaeocurrent reconstructions.

Availability of new Bayesian-delimited gecko names and the importance of character-based species descriptions

Leache & Fujita [[1][1]] present an empirical example of Bayesian species delimitation (BSD; [[2][2]]) to recognize three new species of African geckos from within the range of the widespread taxon Hemidactylus fasciatus , Gray 1842. As with any new method, BSD will undoubtedly generate questions

Determination of genetic diversity within the insular lizard Podarcis tiliguerta using mtDNA sequence data, with a reassessment of the phylogeny of Podarcis

Despite being the predominant reptile group in Southern Europe, the taxonomy of Podarcis Wall lizards is both complex and unstable. Recent attempts to estimate a phylogeny for the genus using molecular methods have been largely unsuccessful, with many poorly resolved nodes and widely different estimates from different studies (Harris and Arnold, 1999; Oliverio et al., 2000; Poulakakis et al., 2003). One possible reason for this is that presently accepted forms may well be species complexes — Podarcis hispanica∗ (Steindachner, 1879) contains several highly genetically distinct lineages all of which may deserve species status based on mitochondrial (Harris and Sa-Sousa, 2002; Harris et al., 2002) and protein electrophoretic data (Pinho et al., 2003). Podarcis erhardii (Bedriaga, 1882) is also probably a species complex (Poulakakis et al., 2003), Podarcis sicula and Podarcis melisellensis contain considerable genetic diversity (Podnar et al., 2004, 2005) although Podarcis lilfordi (Günter,

A hybrid phylogenetic-phylogenomic approach for species tree estimation in African Agama lizards with applications to biogeography, character evolution, and diversification

Africa is renowned for its biodiversity and endemicity, yet little is known about the factors shaping them across the continent. African Agama lizards (45 species) have a pan-continental distribution, making them an ideal model for investigating biogeography. Many species have evolved conspicuous sexually dimorphic traits, including extravagant breeding coloration in adult males, large adult male body sizes, and variability in social systems among colorful versus drab species. We present a comprehensive time-calibrated species tree for Agama, and their close relatives, using a hybrid phylogenetic-phylogenomic approach that combines traditional Sanger sequence data from five loci for 57 species (146 samples) with anchored phylogenomic data from 215 nuclear genes for 23 species. The Sanger data are analyzed using coalescent-based species tree inference using (*)BEAST, and the resulting posterior distribution of species trees is attenuated using the phylogenomic tree as a backbone constraint. The result is a time-calibrated species tree for Agama that includes 95% of all species, multiple samples for most species, strong support for the major clades, and strong support for most of the initial divergence events. Diversification within Agama began approximately 23 million years ago (Ma), and separate radiations in Southern, East, West, and Northern Africa have been diversifying for >10Myr. A suite of traits (morphological, coloration, and sociality) are tightly correlated and show a strong signal of high morphological disparity within clades, whereby the subsequent evolution of convergent phenotypes has accompanied diversification into new biogeographic areas.

Divergence in mitochondrial DNA of Near Eastern water frogs with special reference to the systematic status of Cypriote and Anatolian populations (Anura, Ranidae)

Water frogs from Anatolia, Syria, Jordan, and central Asia (Turkmenistan, Uzbekistan, Kazakhstan) were compared on the basis of the complete mitochondrial (mt) ND3 gene (340 bp), two flanking mt t-RNA gene fragments (26 bp), and a 374 bp fragment of the mt 12S rRNA gene. A total of 27 haplotypes were found among the investigated individuals. Anatolian water frogs differed from Syrian and Jordanian Rana bedriagae by 2.2-3.4% of the analysed sites. The observed divergence (2.8-4.1%) between the Cypriote water frogs and frogs from the surrounding mainland (southern Turkey, west Syria) was in the same range as between R. bedriagae and European R. ridibunda (3.1-3.9%). These results suggest that neither the Cypriote nor the Anatolian water frogs represent R. bedriagae . Furthermore, maximum parsimony and neighbour-joining trees showed a clear subdivision of Asian water frogs into three Anatolian lineages, two central Asian lineages, a Cypriote lineage and the bedriagae lineage. In all trees the Cypriote lineage branches off first and a clade formed by two Anatolian lineages is placed as the sister group of water frogs from Turkmenistan and Uzbekistan, whereas the phylogenetic positions of R. bedriagae , the Ceyhan lineage, the Kazakhstan lineage and R. ridibunda remain unclear.

Cryptic Speciation Patterns in Iranian Rock Lizards Uncovered by Integrative Taxonomy

While traditionally species recognition has been based solely on morphological differences either typological or quantitative, several newly developed methods can be used for a more objective and integrative approach on species delimitation. This may be especially relevant when dealing with cryptic species or species complexes, where high overall resemblance between species is coupled with comparatively high morphological variation within populations. Rock lizards, genus Darevskia, are such an example, as many of its members offer few diagnostic morphological features. Herein, we use a combination of genetic, morphological and ecological criteria to delimit cryptic species within two species complexes, D. chlorogaster and D. defilippii, both distributed in northern Iran. Our analyses are based on molecular information from two nuclear and two mitochondrial genes, morphological data (15 morphometric, 16 meristic and four categorical characters) and eleven newly calculated spatial environmental predictors. The phylogeny inferred for Darevskia confirmed monophyly of each species complex, with each of them comprising several highly divergent clades, especially when compared to other congeners. We identified seven candidate species within each complex, of which three and four species were supported by Bayesian species delimitation within D. chlorogaster and D. defilippii, respectively. Trained with genetically determined clades, Ecological Niche Modeling provided additional support for these cryptic species. Especially those within the D. defilippii-complex exhibit well-differentiated niches. Due to overall morphological resemblance, in a first approach PCA with mixed variables only showed the separation between the two complexes. However, MANCOVA and subsequent Discriminant Analysis performed separately for both complexes allowed for distinction of the species when sample size was large enough, namely within the D. chlorogaster-complex. In conclusion, the results support four new species, which are described herein.

Mitochondrial phylogeography, contact zones and taxonomy of grass snakes (Natrix natrix, N. megalocephala)

Grass snakes (Natrix natrix) represent one of the most widely distributed snake species of the Palaearctic region, ranging from the North African Maghreb region and the Iberian Peninsula through most of Europe and western Asia eastward to the region of Lake Baikal in Central Asia. Within N. natrix, up to 14 distinct subspecies are regarded as valid. In addition, some authors recognize big‐headed grass snakes from western Transcaucasia as a distinct species, N. megalocephala. Based on phylogenetic analyses of a 1984‐bp‐long alignment of mtDNA sequences (ND4+tRNAs, cyt b) of 410 grass snakes, a nearly range‐wide phylogeography is presented for both species. Within N. natrix, 16 terminal mitochondrial clades were identified, most of which conflict with morphologically defined subspecies. These 16 clades correspond to three more inclusive clades from (i) the Iberian Peninsula plus North Africa, (ii) East Europe and Asia and (iii) West Europe including Corso‐Sardinia, the Apennine Peninsula and Sicily. Hypotheses regarding glacial refugia and postglacial range expansions are presented. Refugia were most likely located in each of the southern European peninsulas, Corso‐Sardinia, North Africa, Anatolia and the neighbouring Near and Middle East, where the greatest extant genetic diversity occurs. Multiple distinct microrefugia are inferred for continental Italy plus Sicily, the Balkan Peninsula, Anatolia and the Near and Middle East. Holocene range expansions led to the colonization of more northerly regions and the formation of secondary contact zones. Western Europe was invaded from a refuge within southern France, while Central Europe was reached by two distinct range expansions from the Balkan Peninsula. In Central Europe, there are two contact zones of three distinct mitochondrial clades, and one of these contact zones was theretofore completely unknown. Another contact zone is hypothesized for Eastern Europe, which was colonized, like north‐western Asia, from the Caucasus region. Further contact zones were identified for southern Italy, the Balkans and Transcaucasia. In agreement with previous studies using morphological characters and allozymes, there is no evidence for the distinctiveness of N. megalocephala. Therefore, N. megalocephala is synonymized with N. natrix.

An Early Eocene gecko from Baltic amber and its implications for the evolution of gecko adhesion

A new genus and species of gecko from the Lower Eocene of north-western Russia is described from a superbly preserved specimen in Baltic amber. It is the oldest gekkonid lizard to be represented by more than fragmentary skeletal remains. The digits of the specimen are mostly intact and reveal a unique combination of characters not seen in any living form. Expanded sub-digital scansors on the toes, however, are essentially similar to those of modern climbing geckos and verify the existence of a complex adhesive system 20–30 million years earlier than supported by previously discovered fossil geckos.

The biogeographical assignment of a west Kenyan rain forest remnant: further evidence from analysis of its reptile fauna

Abstract Aim  The Kakamega Forest, western Kenya, has been biogeographically assigned to both lowland and montane forest biomes, or has even been considered to be unique. Most frequently it has been linked with the Guineo‐Congolian rain forest block. The present paper aims to test six alternative hypotheses of the zoogeographical relationships between this forest remnant and other African forests using reptiles as a model group. Reptiles are relatively slow dispersers, compared with flying organisms (Aves and Odonata) on which former hypotheses have been based, and may thus result in a more conservative biogeographical analysis. Location  Kakamega Forest, Kenya, Sub‐Saharan Africa. Methods  The reptile diversity of Kakamega Forest was evaluated by field surveys and data from literature resources. Faunal comparisons of Kakamega Forest with 16 other African forests were conducted by the use of the ‘coefficient of biogeographic resemblance’ using the reptile communities as zoogeographic indicators. Parsimony Analysis of Endemism and Neighbour Joining Analysis of Endemism were used to generate relationship trees based on an occurrence matrix with paup*. Results  The analysis clearly supports the hypothesis that the Kakamega Forest is the easternmost fragment of the Guineo‐Congolian rain forest belt, and thus more closely related to the forests of that Central–West African complex than to any forest further east, such as the Kenyan coastal forests. Many Kenyan reptile species occur exclusively in the Kakamega Forest and its associated forest fragments. Main conclusions  The Kakamega Forest is the only remnant of the Guineo‐Congolian rain forest in the general area. We assume that the low degree of resemblance identified for the Guineo‐Congolian forest and the East African coastal forest reflect the long history of isolation of the two forest types from each other. Kenyan coastal forests may have been historically connected through forest ‘bridges’ of the southern highlands with the Congo forest belt, allowing reptile species to migrate between them. The probability of a second ‘bridge’ located in the region of southern Tanzanian inselbergs is discussed. Although not particularly rich in reptile species, the area should be considered of high national priority for conservation measures.