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The species inventory of global biodiversity is constantly revised and refined by taxonomic research, through the addition of newly discovered species. This almost three century old project provide a knowledge foundation essential for humankind, and notably to develop appropriate conservation strategies. This task relies on the study of millions of specimens housed all around the world in natural history collections. Since two decades, taxonomy generates a plethoric amount of numeric data every year, and notably through the digitization of collection specimens, gradually transforms into a big data science. In this line, the French National Museum of Natural History (MNHN) has embarked into a major research and engineering challenge within its information system, in order to facilitate the transition towards cyber-taxonomic practices which require a facilitated access to data on reference collection specimens housed all over the world. To this end, a first mandatory step is to automatically complete classification data usually associated to collection specimens found in multiple databases. We use here fuzzy approaches to connect one database with other databases and match identical specimens found in each databases together.

Hybridization between the European smooth and palmate newts has recurrently been mentioned in the literature. The only two studies that attempted to quantify the frequency of hybridization and gene admixture between these two species came to strikingly opposite conclusions. According to Arntzen et al (1998, 42 allozymes), hybrids are rare in nature and introgression negligible, while according to Johanet et al (2011, 6 microsatellites), introgressive hybridization is significant and widespread across the shared distribution range. To clarify this question, we implemented high-throughput SNP genotyping with diagnostic biallelic SNPs on 965 specimens sampled across Europe. Our results are in line with Arntzen et al, since only two F1 hybrids were identified in two distinct French localities, and no further hybrid generations or backcrosses. Moreover, reanalysis of 78 of the samples previously studied by Johanet et al. (2011) using our SNPs panel could not reproduce their results, suggesting that microsatellite-based inference overestimated the hybridization frequency between these two species. Since we did not detect methodological issues with the analyses of Johanet et al., our results suggest that SNP approaches outperform microsatellite-based assessments of hybridization frequency, and that conclusions previously published on this topic with a small number of microsatellite loci should be taken with caution, and ideally be repeated with an increased genomic coverage.

AimDiscontinuity in the distribution of genetic diversity (often based on mtDNA) is usually interpreted as evidence for phylogeographic breaks, underlying vicariant units. However, a misleading signal of phylogeographic break can arise in the absence of barrier to gene flow, under mechanisms of isolation by distance (IBD). How and under which conditions phylogeographic breaks can be reliably differentiated from populations evolving under IBD remain unclear. Here, we use multi-locus sequence data from a widely distributed lizard species to address these questions in an empirical setting.LocationMoroccoTaxonSpiny-footed lizard (Acanthodactylus erythrurus), Squamata: LacertidaeMethodsUsing 325 samples from 40 localities, we identified genetic discontinuities withinA. erythrurusbased on a mitochondrial fragment and nine nuclear markers independently. Using the nuclear markers, we then applied linear regression models to investigate whether genetic divergence could be explained by geographical distances alone, or barriers to gene flow (real phylogeographic breaks).ResultA. erythrurusis characterized by an important mitochondrial diversity, with 11 strongly supported phylogeographic lineages with a crown age of 6 Mya. Nuclear markers, however, yielded weak phylogenetic support for these lineages. Using clustering methods based on genotypes at nine nuclear loci, we identified phylogeographic clusters that were partly discordant with the mtDNA lineages. Tests of IBD delimited at least four groups of populations separated by barriers to gene flow, but unambiguous separation of vicariance from IBD remained challenging in several cases.Main conclusionsThe genetic diversity ofA. erythrurusoriginates from a mix of IBD and vicariance, which were difficult to distinguish, and resulted in similar levels of mitochondrial differentiation. These results highlight that phylogeographic breaks inferred from mitochondrial data should be further investigated using multi-locus data and explicit testing to rule out alternative processes generating discontinuities in mitochondrial diversity, including IBD. We identified four groups of populations withinA. erythrurus, separated by barriers to gene flow, but even using nine independent nuclear makers the power of our approach was limited, and further investigation using genome-wide data will be required to resolve the phylogeographic history of this species.

Linnaean and Wallacean shortfalls (Uncertainties on species taxonomy and distribution, respectively) are major factors hampering efficient conservation planning in the current context of biodiversity erosion. These shortfalls concern even widespread and abundant species in relatively well-studied regions such as the Mediterranean biodiversity hotspot which still hosts a large fraction of unrecognised biodiversity, notably in small vertebrates. Species delimitations have long been based on phylogenetic analyses of a small number of standard markers, but accurate lineage identification in this context can be obscured by incomplete lineage sorting, introgression or isolation by distance. Recently, integrative approaches coupling various sets of characters or analyses of contact zones aiming at estimating reproductive isolation (RI) have been advocated instead. Analyses of introgression patterns in contact zone with genomic data represent a powerful way to confirm the existence of independent lineages and estimate the strength of their RI at the same time. The Spiny-footed Lizard Acanthodactylus erythrurus (Schinz, 1833) is widespread in the Iberian Peninsula and the Maghreb and exhibits a large amount of genetic diversity, although the precise number and distribution of its genetic lineages remain poorly understood. We applied a RADseq approach to obtain a genome wide SNPs dataset on a contact zone in central Morocco between the previously described Rif and Middle-Atlas lineages. We show that these two lineages exhibit strong RI across this contact zone, as shown by the limited amount and restricted spatial extant of gene flow. We interpret these results as evidence for species-level divergence of these two lineages. Our study confirms the usefulness of RADseq approaches applied on contact zones for cryptic diversity studies and therefore to resolve Linnaean and Wallacean shortfalls.

The Lygodactylus madagascariensis species group, constituting the subgenus Domerguella, currently contains five valid species of inconspicuous dwarf geckos from Madagascar’s humid forests, but at least 18 deep genetic lineages have been revealed by recent molecular studies. Given the high morphological similarity of these lineages, taxonomic resolution of this astonishing diversity requires efforts to correctly delimit species, as well as assigning the available nomina to the species-level lineages identified. We here combine DNA sequences of one mitochondrial and two nuclear-encoded gene fragments with morphometric measurements and scale counts, and report evidence for a species status of most of the previously identified lineages. In particular, we rely on sympatric and often even syntopic occurrence of several of these lineages without evidence for genetic admixture, and consistent with subtle morphological differences. Furthermore, the very high divergences of 7.4–23.8% pairwise distances in the relatively conserved mitochondrial 16S rRNA gene, combined with a lack of haplotype sharing in the nuclear-encoded genes and differences in scale counts convinced us that most of the other, allopatrically distributed lineages also represent distinct species. We elevate L. madagascariensis petteri to species level and formally name eight new species: L. salvi sp. nov., a species from the Sambirano region in northern Madagascar, previously called L. sp. 8; L. tantsaha sp. nov. (L. sp. 10), a species occurring sympatrically with L. madagascariensis and L. petteri on Montagne d’Ambre in far northern Madagascar; L. roellae sp. nov. (L. sp. 17), a species characterized by a striped coloration in all known specimens, from northern Madagascar; L. winki sp. nov. (L. sp. 18), an unstriped species from northern Madagascar but belonging to a subclade mostly distributed in the eastern rainforests of the island; L. ulli sp. nov. (L. sp. 21), a species from the same subclade as L. winki but known only from the Marojejy Massif in the North East; L. fritzi sp. nov. (L. sp. 11), a further species of this subclade from coastal lowlands in the Northern Central East; L. hodikazo sp. nov. (L. sp. 23) known from a single specimen collected at the Tsingy de Bemaraha and therefore the only Domerguella species known from the West region of Madagascar; and L. hapei sp. nov. (L. sp. 26), an enigmatic species from the Sambirano region characterized by a striped pattern on the throat that is otherwise unknown in the subgenus. Three additional deep mitochondrial lineages of Domerguella were identified in our analysis, but could not be further analyzed due to the lack or scarcity of voucher specimens. More field work and collection of voucher specimens is needed to understand their status. Furthermore, the taxonomy of the Domerguella subclade occurring in eastern Madagascar, with three described species (L. guibei, L. miops, L. fritzi), two synonyms (L. septemtuberculatus, Microscalabotes spinulifer) and at least two further deep genetic lineages co-occurring in a relatively small area, requires further revisionary work, possibly aided by target-enrichment sequencing of the respective name-bearing types.

The Paroedura bastardi clade, a subgroup of the Madagascan gecko genus Paroedura, currently comprises four nominal species: P. bastardi, supposedly widely distributed in southern and western Madagascar, P. ibityensis, a montane endemic, and P. tanjaka and P. neglecta, both restricted to the central west region of the island. Previous work has shown that Paroedura bastardi is a species complex with several strongly divergent mitochondrial lineages. Based on one mitochondrial and two nuclear markers, plus detailed morphological data, we undertake an integrative revision of this species complex. Using a representative sampling for seven nuclear and five mitochondrial genes we furthermore propose a phylogenetic hypothesis of relationships among the species in this clade. Our analyses reveal at least three distinct and independent evolutionary lineages currently referred to P. bastardi. Conclusive evidence for the species status of these lineages comes from multiple cases of syntopic occurrence without genetic admixture or morphological intermediates, suggesting reproductive isolation. We discuss the relevance of this line of evidence and the conditions under which concordant differentiation in unlinked loci under sympatry provides a powerful approach to species delimitation, and taxonomically implement our findings by (1) designating a lectotype for Paroedura bastardi, now restricted to the extreme South-East of Madagascar, (2) resurrecting of the binomen Paroedura guibeae Dixon & Kroll, 1974, which is applied to the species predominantly distributed in the South-West, and (3) describing a third species, Paroedura rennerae sp. nov., which has the northernmost distribution within the species complex.

Research data are available on the Mendeley Data platform (https://doi.org/10.17632/xfx2yhw949.2), which include datasets, input files and R scripts of the analyses., Appendix A. Supplementary material Supplementary data to this article can be found online at https://doi. org/10.1016/j.ympev.2022.107635, Most of the unique and diverse vertebrate fauna that inhabits Madagascar derives from in situ diversification from colonisers that reached this continental island through overseas dispersal. The endemic Malagasy Scincinae lizards are amongst the most species-rich squamate groups on the island. They colonised all bioclimatic zones and display many ecomorphological adaptations to a fossorial (burrowing) lifestyle. Here we propose a new phylogenetic hypothesis for their diversification based on the largest taxon sampling so far compiled for this group. We estimated divergence times and investigated several aspects of their diversification (diversification rate, body size and fossorial lifestyle evolution, and biogeography). We found that diversification rate was constant throughout most of the evolutionary history of the group, but decreased over the last 6–4 million years and independently from body size and fossorial lifestyle evolution. Fossoriality has evolved from fully quadrupedal ancestors at least five times independently, which demonstrates that even complex morphological syndromes – in this case involving traits such as limb regression, body elongation, modification of cephalic scalation, depigmentation, and eyes and ear-opening regression – can evolve repeatedly and independently given enough time and eco-evolutionary advantages. Initial diversification of the group likely occurred in forests, and the divergence of sand-swimmer genera around 20 Ma appears linked to a period of aridification. Our results show that the large phenotypic variability of Malagasy Scincinae has not influenced diversification rate and that their rich species diversity results from a constant accumulation of lineages through time. By compiling large geographic and trait-related datasets together with the computation of a new time tree for the group, our study contributes important insights on the diversification of Malagasy vertebrates., Portuguese National Funds through FCT (Fundação para a Ciência e a Tecnologia) supported this work with the projects UIDP/50027/2020 and PTDC/BIA-EVL/31254/2017, Contract grant to AC [2020.00823.CEECIND/CP1601/CT0003, PD/BD/128493/2017, SFRH/BD/102495/2014, SFRH/BPD/111015/2015, Agencia Española de Investigación, with a Juan de la Cierva-Formación grant [FCI2019-039443-I/AEI/10.13039/501100011033], Swedish Research Council [2019-05191], Swiss National Science Foundation [PCEFP3_187012], Swedish Research Council [VR: 2019-04739]

International audience; A wide range of data types can be used to delimit species and various computer-based tools dedicated to this task are now available. Although these formalized approaches have significantly contributed to increase the objectivity of species delimitation (SD) under different assumptions, they are not routinely used by alpha-taxonomists. One obvious shortcoming is the lack of interoperability among the various independently developed SD programs. Given the frequent incongruences between species partitions inferred by different SD approaches, researchers applying these methods often seek to compare these alternative species partitions to evaluate the robustness of the species boundaries. This procedure is excessively time consuming at present, and the lack of a standard format for species partitions is a major obstacle. Here, we propose a standardized format, SPART, to enable compatibility between different SD tools exporting or importing partitions. This format reports the partitions and describes, for each of them, the assignment of individuals to the “inferred species”. The syntax also allows support values to be optionally reported, as well as original trees and the full command lines used in the respective SD analyses. Two variants of this format are proposed, overall using the same terminology but presenting the data either optimized for human readability (matricial SPART) or in a format in which each partition forms a separate block (SPART.XML). ABGD, DELINEATE, GMYC, PTP and TR2 have already been adapted to output SPART files and a new version of LIMES has been developed to import, export, merge and split them.

Our knowledge of the conservation status of reptiles, the most diverse class of terrestrial vertebrates, has improved dramatically over the past decade, but still lags behind that of the other tetrapod groups. Here, we conduct the first comprehensive evaluation (~92% of the world’s ~1,714 described species) of the conservation status of skinks (Scincidae), a speciose reptile family with a worldwide distribution. Using International Union for Conservation of Nature (IUCN) criteria, we report that ~20% of species are threatened with extinction, and nine species are Extinct or Extinct in the Wild. The highest levels of threat are evident in Madagascar and the Neotropics, and in the subfamilies Mabuyinae, Eugongylinae and Scincinae. The vast majority of threatened skink species were listed based primarily on their small geographic ranges (Criterion B, 83%; Criterion D2, 13%). Although the population trend of 42% of species was stable, 14% have declining populations. The key threats to skinks are habitat loss due to agriculture, invasive species, and biological resource use (e.g., hunting, timber harvesting). The distributions of 61% of species do not overlap with protected areas. Despite our improved knowledge of the conservation status of the world’s skinks, 8% of species remain to be assessed, and 14% are listed as Data Deficient. The conservation status of almost a quarter of the world’s skink species thus remains unknown. We use our updated knowledge of the conservation status of the group to develop and outline the priorities for the conservation assessment and management of the world’s skink species.

International audience; Burrowing is a common behavior in vertebrates. An underground life-style offers many advantages but also poses important challenges including the high energetic cost of burrowing. Scolecophidians are a group of morphologically derived subterranean snakes that show great diversity in form and function. Although it has been suggested that leptotyphlopids and anomalepidids mostly use existing underground passageways, typhlopids are thought to create their own burrows. However, the mechanisms used to create burrows and the associated forces that animals may be able to generate remain unknown. Here, we provide the first data on push forces in scolecophidians and compare them with those in some burrowing alethinophidian snakes. Our results show that typhlopids are capable of generating higher forces for a given size than other snakes. The observed differences are not due to variation in body diameter or length, suggesting fundamental differences in the mechanics of burrowing or the way in which axial muscles are used. Qualitative observations of skull and vertebral shape suggest that the higher forces exerted by typhlopids may have impacted the evolution of their anatomy. Our results provide the basis for future studies exploring the diversity of form and function in this fascinating group of animals. Quantitative comparisons of the cranial and vertebral shape in addition to collecting functional and ecological data on a wider array of species would be particularly important to test the patterns described here.

While powerful and user-friendly software suites exist for phylogenetics, and an impressive cybertaxomic infrastructure of online species databases has been set up in the past two decades, software targeted explicitly at facilitating alpha-taxonomic work, i.e., delimiting and diagnosing species, is still in its infancy. Here we present a project to develop a bioinformatic toolkit for taxonomy, based on open-source Python code, including tools focusing on species delimitation and diagnosis and centered around specimen identifiers. At the core of iTaxoTools is user-friendliness, with numerous autocorrect options for data files and with intuitive graphical user interfaces. Assembled standalone executables for all tools or a suite of tools with a launcher window will be distributed for Windows, Linux, and Mac OS systems, and in the future also implemented on a web server. The initial version (iTaxoTools 0.1) distributed with this paper (https://github.com/iTaxoTools/iTaxoTools-Executables) contains graphical user interface (GUI) versions of six species delimitation programs (ABGD, ASAP, DELINEATE, GMYC, PTP, tr2) and a simple threshold-clustering delimitation tool. There are also new Python implementations of existing algorithms, including tools to compute pairwise DNA distances, ultrametric time trees based on non-parametric rate smoothing, species-diagnostic nucleotide positions, and standard morphometric analyses. Other utilities convert among different formats of molecular sequences, geographical coordinates, and units; merge, split and prune sequence files, tables and species partition files; and perform simple statistical tests. As a future perspective, we envisage iTaxoTools to become part of a bioinformatic pipeline for next-generation taxonomy that accelerates the inventory of life while maintaining high-quality species hypotheses. The open source code and binaries of all tools are available from Github (https://github.com/iTaxoTools) and further information from the website (http://itaxotools.org).

The genus Scaphiophryne (Anura: Microhylidae) contains at least nine species that, together with their probable sister genus Paradoxophyla, form the Madagascar-endemic subfamily Scaphiophryninae. Scaphiophryne are robust burrowing frogs with explosive breeding behavior and are characterized by a unique larval morphology, being intermediate between that of filter-feeding and generalized tadpole types. Based on tadpole morphology, the nominal subgenus Scaphiophryne can be distinguished from the subgenus Pseudohemisus, a clade occurring in arid biomes with a largely unsolved taxonomy and nomenclature. Here we combine a comprehensive molecular phylogeny of all scaphiophrynines based on five mitochondrial and two nuclear genes with a re-examination of the name-bearing type specimens of the seven species-level nomina assignable to the subgenus Pseudohemisus. Our molecular phylogeny supports monophyly of the two subgenera and assigns the enigmatic S. matsoko to the subgenus Scaphiophryne. In this subgenus, our analysis confirms all currently accepted species as independent evolutionary lineages based on concordant differentiation in mitochondrial and nuclear genes, and provides evidence for a combination of low mitochondrial divergence, distinct nuclear divergence, and high nuclear heterozygosity. Yet, contrary to previous hypotheses, we do not find evidence of ongoing hybridization. In the subgenus Pseudohemisus we find support for three clearly delimited species-level lineages, one of which contains two deep conspecific lineages that warrant further study. To stabilize the taxonomy in this subgenus, and considering the very poor state of preservation of several type series and the heterogeneity of some of them, we here designate lectotypes for the nomina: Hemisus obscurus Grandidier, 1872; Calophrynus calcaratus Mocquard, 1895; Pseudohemisus verrucosus Angel, 1930; Pseudohemisus longimanus Angel, 1930; and Pseudohemisus longimanus var. melanopleura Angel, 1934. Taxonomically, we accept Scaphiophryne brevis (Boulenger, 1896) as a well-defined species; restrict S. calcarata (Mocquard, 1995) to the southern lineage of the S. calcarata complex; resurrect Hemisus obscurus Grandidier, 1872 (previously considered to be a dubious name or nomen inquirendum) as Scaphiophryne obscura to refer to the western and north-western lineage of the S. calcarata complex; consider Pseudohemisus verrucosus Angel, 1930 as junior synonym of Scaphiophryne brevis; and consider Pseudohemisus longimanus Angel, 1930 and Pseudohemisus longimanus var. melanopleura Angel, 1934 as junior synonyms of Scaphiophryne obscura. Finally, to accelerate taxonomic progress, we (i) outline a set of criteria by which such old and ambiguous nomina can be applied to known lineages, (ii) suggest that the International Code of Zoological Nomenclature should slightly broaden the conditions under which neotypes are to be designated, and (iii) propose to redouble discussions on the introduction of the concept of the epitype to zoology.

Natural history collections are leading successful large-scale projects of specimen digitization (images, metadata, DNA barcodes), thereby transforming taxonomy into a big data science. Yet, little effort has been directed towards safeguarding and subsequently mobilizing the considerable amount of original data generated during the process of naming 15,000–20,000 species every year. From the perspective of alpha-taxonomists, we provide a review of the properties and diversity of taxonomic data, assess their volume and use, and establish criteria for optimizing data repositories. We surveyed 4113 alpha-taxonomic studies in representative journals for 2002, 2010, and 2018, and found an increasing yet comparatively limited use of molecular data in species diagnosis and description. In 2018, of the 2661 papers published in specialized taxonomic journals, molecular data were widely used in mycology (94%), regularly in vertebrates (53%), but rarely in botany (15%) and entomology (10%). Images play an important role in taxonomic research on all taxa, with photographs used in >80% and drawings in 58% of the surveyed papers. The use of omics (high-throughput) approaches or 3D documentation is still rare. Improved archiving strategies for metabarcoding consensus reads, genome and transcriptome assemblies, and chemical and metabolomic data could help to mobilize the wealth of high-throughput data for alpha-taxonomy. Because long-term—ideally perpetual—data storage is of particular importance for taxonomy, energy footprint reduction via less storage-demanding formats is a priority if their information content suffices for the purpose of taxonomic studies. Whereas taxonomic assignments are quasifacts for most biological disciplines, they remain hypotheses pertaining to evolutionary relatedness of individuals for alpha-taxonomy. For this reason, an improved reuse of taxonomic data, including machine-learning-based species identification and delimitation pipelines, requires a cyberspecimen approach—linking data via unique specimen identifiers, and thereby making them findable, accessible, interoperable, and reusable for taxonomic research. This poses both qualitative challenges to adapt the existing infrastructure of data centers to a specimen-centered concept and quantitative challenges to host and connect an estimated \documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{upgreek} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} }{}$ \le $\end{document}2 million images produced per year by alpha-taxonomic studies, plus many millions of images from digitization campaigns. Of the 30,000–40,000 taxonomists globally, many are thought to be nonprofessionals, and capturing the data for online storage and reuse therefore requires low-complexity submission workflows and cost-free repository use. Expert taxonomists are the main stakeholders able to identify and formalize the needs of the discipline; their expertise is needed to implement the envisioned virtual collections of cyberspecimens. [Big data; cyberspecimen; new species; omics; repositories; specimen identifier; taxonomy; taxonomic data.]

The ability to rapidly generate and share molecular, visual, and acoustic data, and to compare them with existing information, and thereby to detect and name biological entities is fundamentally changing our understanding of evolutionary relationships among organisms and is also impacting taxonomy. Harnessing taxonomic data for rapid, automated species identification by machine learning tools or DNA metabarcoding techniques has great potential but will require their review, accessible storage, comprehensive comparison, and integration with prior knowledge and information. Currently, data production, management, and sharing in taxonomic studies are not keeping pace with these needs. Indeed, a survey of recent taxonomic publications provides evidence that few species descriptions in zoology and botany incorporate DNA sequence data. The use of modern high-throughput (-omics) data is so far the exception in alpha-taxonomy, although they are easily stored in GenBank and similar databases. By contrast, for the more routinely used image data, the problem is that they are rarely made available in openly accessible repositories. Improved sharing and re-using of both types of data requires institutions that maintain long-term data storage and capacity with workable, user-friendly but highly automated pipelines. Top priority should be given to standardization and pipeline development for the easy submission and storage of machine-readable data (e.g., images, audio files, videos, tables of measurements). The taxonomic community in Germany and the German Federation for Biological Data are researching options for a higher level of automation, improved linking among data submission and storage platforms, and for making existing taxonomic information more readily accessible.

Trade-offs are thought to be important in constraining evolutionary divergence as they may limit phenotypic diversification. The cranial system plays a vital role in many functions including defensive, territorial, predatory and feeding behaviours in addition to housing the brain and sensory systems. Consequently, the morphology of the cranial system is affected by a combination of selective pressures that may induce functional trade-offs. Limbless, head-first burrowers are thought to be constrained in their cranial morphology as narrow heads may provide a functional advantage for burrowing. However, having a wide and large head is likely beneficial in terms of bite performance. We used 15 skink species to test for the existence of trade-offs between maximal push and bite forces, and explored the patterns of covariation between external head and body morphology and performance. Our data show that there is no evidence of a trade-off between bite and burrowing in terms of maximal force. Species that generate high push forces also generate high bite forces. Our data also show that overall head size covaries with both performance traits. However, future studies exploring trade-offs between force and speed or the energetic cost of burrowing may reveal other trade-offs.

International audience; Currently the planet is inhabited by several millions of extremely diversified species. Not all of them arouse emotions of the same nature or intensity in humans. Little is known about the extent of our affective responses toward them and the factors that may explain these differences. Our online survey involved 3500 raters who had to make choices depending on specific questions designed to either assess their empathic perceptions or their compassionate reactions toward an extended photographic sampling of organisms. Results show a strong negative correlation between empathy scores and the divergence time separating them from us. However, beyond a certain time of divergence, our empathic perceptions stabilize at a minimum level. Compassion scores, although based on less spontaneous choices, remain strongly correlated to empathy scores and time of divergence. The mosaic of features characterizing humans has been acquired gradually over the course of the evolution, and the phylogenetically closer a species is to us, the more it shares common traits with us. Our results could be explained by the fact that many of these traits may arouse sensory biases. These anthropomorphic signals could be able to mobilize cognitive circuitry and to trigger prosocial behaviors usually at work in human relationships. "Sympathy beyond the confines of man, that is, humanity to the lower animals, seems to be one of the latest moral acquisitions. (…). This virtue, one of the noblest with which man is endowed, seems to arise incidentally from our sympathies becoming more tender and more widely diffused, until they are extended to all sentient beings". Charles Darwin, 1871

Background: GLORIA-AF (Global Registry on Long-Term Oral Antithrombotic Treatment in Patients with Atrial Fibrillation) is a prospective, global registry program describing antithrombotic t...

The phylogenetic relationships between the three main clades of worm snakes remain controversial. This question is, however, crucial to elucidate the origin of the successful snake radiation, as these burrowing and miniaturized wormlike organisms represent the earliest branching clades within the snake tree. The present molecular phylogenetic study, intended to minimize the amount of missing data, provides fully resolved inter-subfamilial relationships among Typhlopidae. It also brings robust evidence that worm snakes (Scolecophidia) are paraphyletic, with the scolecophidian family Anomalepididae recovered with strong support as sister clade to the 'typical snakes' (Alethinophidia). Ancestral state reconstructions applied to three different traits strongly associated to a burrowing life-style (scolecoidy, absence of retinal cones and microstomy) provide results in favour of a burrowing origin of snakes, and suggest that worm snakes might be the only extant fossorial representatives of the primordial snake incursion towards an underground environment.

Motivation Species delimitation (SD) is on the verge of becoming a fully fledged research field in systematics, but the variety of available approaches tends to result in significant—sometimes striking—incongruences, when tested comparatively with a given taxonomic sampling. Results We present LIMES, an automatic calculation tool which qualitatively compares species partitions obtained by distinct SD approaches, regardless of their respective theoretical backgrounds, and even in absence of reference topology. The program implements four different previously published indexes, and allows their automated calculation. Availability and implementation LIMES is freely downloadable at www.limes.cnrs.fr. Supplementary information Supplementary data are available at Bioinformatics online.

The family Xenophidiidae is an extremely rare and little-known family of snakes that contains only two species, Xenophidion acanthognathus Günther & Manthey and X. schaeferi Günther & Manthey that are each known only from their holotypes. We report on the rediscovery of X. schaeferi from two new locations in southern Peninsular Malaysia. The new specimens differ markedly from the holotype of X. schaeferi in colour pattern and are more akin to the patterning of X. acanthognathus from Borneo. However, molecular analyses of one of the new specimens reveal that it only differs from the holotype of X. schaeferi by a minimum sequence divergence of 0.27%. An expanded description of the species based on these specimens is provided and phenotypic variation within the species is discussed. We also report the first record of the genus Xenophidion from West Sumatra, Indonesia. These sensational discoveries continue to underscore the fact that southern Peninsular Malaysia and upland areas of Southeast Asia in general still harbour unrealized herpetological diversity and are still in need of field research-based taxonomic studies.

Syncytins are envelope genes from endogenous retroviruses that have been captured during evolution for a function in placentation. They have been found in all placental mammals in which they have been searched, including marsupials. Placental structures are not restricted to mammals but also emerged in some other vertebrates, most frequently in lizards, such as the viviparous

Syncytins are envelope genes from endogenous retroviruses that have been captured during evolution for a function in placentation. They have been found in all placental mammals in which they have been searched, including marsupials. Placental structures are not restricted to mammals but also emerged in some other vertebrates, most frequently in lizards, such as the viviparous Mabuya Scincidae. Here, we performed high-throughput RNA sequencing of a Mabuya placenta transcriptome and screened for the presence of retroviral env genes with a full-length ORF. We identified one such gene, which we named "syncytin-Mab1," that has all the characteristics expected for a syncytin gene. It encodes a membrane-bound envelope protein with fusogenic activity ex vivo, is expressed at the placental level as revealed by in situ hybridization and immunohistochemistry, and is conserved in all Mabuya species tested, spanning over 25 My of evolution. Its cognate receptor, required for its fusogenic activity, was searched for by a screening assay using the GeneBridge4 human/Chinese hamster radiation hybrid panel and found to be the MPZL1 gene, previously identified in mammals as a signal-transducing transmembrane protein involved in cell migration. Together, these results show that syncytin capture is not restricted to placental mammals, but can also take place in the rare nonmammalian vertebrates in which a viviparous placentotrophic mode of reproduction emerged. It suggests that similar molecular tools have been used for the convergent evolution of placentation in independently evolved and highly distant vertebrates.

Novel ventrally positioned extra-cloacal glands were discovered through examination of cloacal histology of female thamnophiine snakes. The glands discovered, termed ventral urodeal glands, are paired and empty into the caudal extremity of the urodeum, lateral to the opening of the urodeal sphincter, through primary ducts that resemble folds of the urodeal wall. In concordance with historical literature, these glands are the first cloacal glands described in snakes that definitively empty into the urodeum from a ventral position. Ventral urodeal glands are complex, with multiple branching ducts that empty the terminal alveolar glands. The epithelial linings of the branching duct networks are stratified (primary and secondary ducts), pseudostratified (tertiary ducts), or simple (quaternary ducts and terminal alveoli). The epithelia of the terminal alveoli stain weakly with hematoxylin, and an eosinophilic secretory material is common in the lumina of alveoli. The secretion produced in the alveoli stains positive with the periodic acid-Schiff's procedure, negative with brilliant blue, and positive with fast green, indicating that the majority of the secretion is composed of a neutral carbohydrate moiety with a minor protein component. Ventral urodeal glands were observed in all taxa of Nerodia examined, except N. cyclopion, and were not observed in any other taxa of Thamnophiini. Through optimization to Thamnophiini topologies, ventral urodeal glands either evolved once on the branch leading to Nerodia and were subsequently lost on the branch leading to N. cyclopion, or they evolved independently on the branches leading to N. floridana and all other species of Nerodia. Like many of the cloacal glands in squamates, the function of ventral urodeal glands is unknown, and further investigation is needed to elucidate the function of the secretions of these glands.

Lizards of the genus Trachylepis are a species-rich group of skinks mainly inhabiting Africa, Madagascar, and several other islands in the western Indian Ocean. All except one probably introduced species of Madagascan Trachylepis are endemic. Two species groups have been distinguished on the basis of subocular scale shape but their phylogenetic relationships remained unclear. We inferred a multilocus phylogeny of the Madagascan Trachylepis species, based on a concatenated dataset of 3261 bp from 3 mitochondrial and 4 nuclear genes with a dense Madagascan taxon sampling and find high support for the monophyly of the endemic Madagascan Trachylepis. The two species groups in Madagascar are highly supported as clades. The highland species T. boettgeri is nested in the T. aureopunctata species group of mainly arid-adapted species, suggesting a colonization of highland swamps by ancestors inhabiting dry western Madagascar. The Seychellois species were sister to the T. maculilabris/T. comorensis clade, suggesting their origin directly out of Africa as with Seychellois chameleons. In Madagascar, a high intraspecific molecular variation was confirmed for T. gravenhorstii, T. elegans, and T. vato, indicating a need for taxonomic revision.

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Among the endemic biota of Madagascar, skinks are a diverse radiation of lizards that exhibit a striking ecomorphological variation, and could provide an interesting system to study body-form evolution in squamate reptiles. We provide a new phylogenetic hypothesis for Malagasy skinks of the subfamily Scincinae based on an extended molecular dataset comprising 8060 bp from three mitochondrial and nine nuclear loci. Our analysis also increases taxon sampling of the genus Amphiglossus by including 16 out of 25 nominal species. Additionally, we examined whether the molecular phylogenetic patterns coincide with morphological differentiation in the species currently assigned to this genus. Various methods of inference recover a mostly strongly supported phylogeny with three main clades of Amphiglossus. However, relationships among these three clades and the limb-reduced genera Grandidierina, Voeltzkowia and Pygomeles remain uncertain. Supported by a variety of morphological differences (predominantly related to the degree of body elongation), but considering the remaining phylogenetic uncertainty, we propose a redefinition of Amphiglossus into three different genera (Amphiglossus sensu stricto, Flexiseps new genus, and Brachyseps new genus) to remove the non-monophyly of Amphiglossus sensu lato and to facilitate future studies on this fascinating group of lizards.

International audience; In a previous study, Miralles and Vences (2013) compared seven different methods of species delimitation applied to the genus Madascincus. While focusing on methodological aspects their study involved an extensive data set of multilocus DNA sequences and of comparative morphology. On this basis they emphasized the need of revising the taxonomy of Madascincus, and revealed the existence of at least two well-supported candidate species. The present paper provides formal descriptions of these two taxa: (1) M. miafina sp. n., a species from dry areas of northern Madagascar, morphologically very similar to M. polleni (although both species are not retrieved as sister taxa), and (2) M. pyrurus sp. n., a montane species occurring >1500 m above sea level, endemic to the central highlands of Madagascar (Ibity and Itremo Massifs). Phylogenetically, M. pyrurus is the sister species of M. igneocaudatus, a taxon restricted to the dry littoral regions of the south and south-west of Madagascar in lowlands

Siegel D.S., Miralles A, Trauth S.E. and Aldridge R.D. 2012. The phylogeneticdistribution and morphological variation of the ‘pouch’ in female snakes. — ActaZoologica (Stockholm) 93: 400–408.The urodaeum of female snakes is a cham ber that receives the oviducts, urinaryducts, and intestine. Previous studies have indicated that a derived region ofunknown function persists between the urodaeum and oviducts in some snakesof the Colubroides. This structure was recently termed the ‘pouch’; however, itis also commonly referred to as the vaginal pouch. A broad sampling of femalesnakes revealed that the pouch evolved once on the branch leading to Colubro-ides. The presence of the pouch in Colubroides is not unambiguous, as this fea-ture was lost in some members of the Atractaspididae and Lamprophiidae.Variation was also observed in pouch morphology between taxa of the Colubro-ides. Variation included the following: (1) whether the pouch was bifurcated andifso,werethearmsofthepouchseparated caudally, (2) whether separatedpouch arms were split by a caudal bifurcation of the urodaeum or through med-ial septa of the pouch arms, and (3) the absence or presence of oviducal papillaeinvading the cranial ⁄lateral extremity of the pouch. The variation in pouch mor-phology observed does not appear to be correlated with the evolutionary historyof Colubroides and polymorphism within families is common.Dustin S. Siegel, Department of Biology, Saint Louis University, 3507 LacledeAvenue, St. Louis, Missouri 63103, USA. E-mail: dsiegel2@slu.eduIntroductionThe female snake cloaca is comprised of a caudal procto-daeum and cranial urodaeum (Siegel et al. 2011a). The uri-nary ducts, intestine, and oviducts communicate with theurodaeum, whereas the proctodaeum communicates theurodaeum with the external environment (Fig. 1). Four typesof cloacae exist in female snakes and are defined by variationof the oviducal ⁄cloacal junction (Siegel et al. 2011b). Type I isfound only in Scolecophidia and is defined by the completeabsence of one oviducal ⁄urodaeal junction. Type II is found insome lizards, Boidae, Loxocemidae, Pythonidae, some Uro-peltidae, and Xenopeltidae and is defined by oviducts passingdorsal or ventralto the urodaeum before a caudalcommunica-tion with the urodaeum. The Type III cloaca is defined by ovi-ducts that insert on the medial horizon of the urodaeum and isfound in some lizards, Caenophidia, and some Uropeltidae.Type IV cloaca possess urodaeal diverticuli that communicatewith the oviducts dorsal to the main urodaeal body, and thiscloaca is restricted to Aniliidae and Tropidophiidae.The ‘pouch’ (or vaginal pouch) is a modified portion of thecranial urodaeum or caudal oviductsinsomefemalesnakes(Siegel et al. 2011a). Giacomini (1893) initially described thepouch in a host of snakes from the Colubroides sensu (Zaheret al. 2009; Caenophidia sans Acrochordidae). Thereafter,multiple authors also noted this distinctive region in othersnakes of the Colubroides (i.e., Elapidae, Colubridae, andDipsadidae; Cope 1898; Ludwig and Rahn 1943; Fox 1956;Gabe and Saint-Girons 1965; Aldridge 1992; Uribe et al.1998; Sa´nchez-Marti´nez et al. 2007; Siegel et al. 2011a), butnot in the more basal lineages of snakes (i.e., Scolecophidiaand Pythonidae; Fox and Dessauer 1962; Gabe and Saint-Girons 1965). Thus, Siegel et al. (2011a) hypothesized thatthe pouch was a synapomorphy for the Colubroides, a group-ing of snakes that has received robust phylogenetic support(e.g., Lawson et al. 2005; Zaher et al. 2009).Whereas the urodaeum is lined by a bistratified-to-stratifiedsquamous ⁄cuboidal ⁄columnar epithelium (Gabe and Saint-Girons 1965; Siegel et al. 2011a), a simple to pseudostratifiedtall columnar epithelium lines the pouch. The majority of the

In squamate reptiles the rostral shield constitutes one of the most advanced cases of reduction in the number of scales in the rostral region, an evolutionary trend clearly associated with a burrowing lifestyle. This structure is characterized by the fusion of the rostral scale with all adjacent scales into a large, smooth and conical plate covering the snout, totally encompassing the nostrils, with a horizontal groove running posteriorly from either nostril. In lizards this structure evolved several times independently, in various lineages of limbless skinks and in the family Dibamidae. We performed a multilocus phylogenetic analysis of combined mitochondrial and nuclear DNA sequences from the fossorial genus Paracontias, including P. vermisaurus, a new species described herein under an integrative taxonomic approach. The resulting phylogeny supports monophyly of Paracontias, with the following internal topology: [P. kankana (P. vermisaurus sp. n. (((P. minimus + P. brocchii) (P. manify + P. hildebrandti)) (P. rothschildi + P. fasika)))]. The molecular data, coupled with a comparative morphological study, allows us to investigate the evolution of the snout scales into a single large rostral shield in Paracontias. We discuss the evolutionary processes through which the rostral shield may have originated (e.g. fusion of scales, number and order of steps involved), and conclude that intuitive and apparently obvious hypotheses for scale homologies based on position and size only (as usually formulated in squamate taxonomy) may be highly misleading, even in closely related species. We develop the hypothesis that the rostral shield may provide several functional advantages for fossorial species in facilitating burrowing and protecting the head from strong physical stress, e.g. smoother surface reducing friction between the tegument and the substrate, reduction in the number of flexible sutures resulting in strengthened tegument, and the rostral tip likely playing a role as a shock-absorbing buffer.

Since the advent of molecular character sets in phylogenetic systematics our understanding of the evolutionary history of snakes has changed considerably. In some cases the novel topologies reconstructed from molecular datasets have left researchers puzzled, as no morphological feature seems to support the new relationships found. This is the case for ‘Amerophidia’sensu Vidal et al. (2007; Biology of the Boas and Pythons, Eagle Mountain: Eagle Mountain Publishing; Aniliidae + Tropidophiidae), a grouping of the Red Pipesnakes and Neotropical Dwarf Boas. We contend that in some cases the apparent lack of historical morphological support for the molecular phylogenies is due to our poor understanding of the organisms as a whole, and not the complete lack of morphological support for controversial clades. For example, we found novel evidence from reproductive anatomy that demonstrates a unique association of the oviducts and cloaca in Amerophidia. Whereas in all other female squamates the oviducts communicate directly with the cloaca, the oviducts of Aniliidae and Tropidophiidae communicate with diverticuli of the cloaca. At present this is the only unambiguous synapomorphy for the Amerophidia. We feel that confirmation of controversial molecular relationships will revolve around the investigation of non-traditional morphological characters such as reproductive anatomy.

Miralles, A., Vasconcelos, R., Perera, A., Harris, D. J. & Carranza, S. (2010). An integrative taxonomic revision of the Cape Verdean skinks (Squamata, Scincidae). —Zoologica Scripta, 40, 16–44. A comprehensive taxonomic revision of the Cape Verdean skinks is proposed based on an integrative approach combining (i) a phylogenetic study pooling all the previously published molecular data, (ii) new population genetic analyses using mitochondrial and nuclear data resulting from additional sampling, together with (iii) a morphological study based on an extensive examination of the scalation and colour patterns of 516 live and museum specimens, including most of the types. All Cape Verdean species of skinks presently recognised, formerly regarded as members of the genera Mabuya Fitzinger, 1826 and Macroscincus Bocage, 1873 are considered as members of the Cape Verdean endemic genus Chioninia Gray, 1845. The new phylogeny and networks obtained are congruent with the previously published phylogenetic studies, although suggesting older colonization events (between 11.6 and 0.8 Myr old), and indicate the need for taxonomic changes. Intraspecific diversity has been analysed and points to a very recent expansion of Chioninia delalandii on the southern islands and its introduction on Maio, to a close connection between Chioninia stangeri island populations due to Pleistocene sea-level falls and to a generally low haplotypic diversity due to the ecological and geological characteristics of the archipelago. Three new consistent morphological synapomorphies supporting two of the four main clades of the genus have been identified. The complex taxonomic status of Euprepes fogoensis O’Shaughnessy, 1874 has been resolved and a lectotype has been designated for this species; Chioninia fogoensis nicolauensis (Schleich, 1987) is elevated to species rank, whereas Chioninia fogoensis antaoensis (Schleich, 1987) is now regarded as a junior subjective synonym of C. fogoensis. Additionally, one new subspecies of Chioninia vaillanti and two of Chioninia spinalis are described (Chioninia vaillanti xanthotis ssp. n., Chioninia spinalis santiagoensis ssp. n. and Chioninia spinalis boavistensis ssp. n.) and a lectotype has been designated for Mabuia spinalis Boulenger, 1906. Finally, an identification key for the Chioninia species is presented.

A specimen of the colubrid genus Ahaetulla Link, 1807 collected in 2002 in Jambi Province, Sumatra, Indonesia, proves to be the first record of Ahaetulla mycterizans (Linnaeus, 1758) for this Indonesian island. This species was previously known from Java, West Malaysia and southern Peninsular Thailand. The discovery of this specimen constitutes an opportunity to redefine and illustrate this rare and poorly known species and to compare it with the more common Ahaetulla prasina (Boie, 1827). Additionally, an identification key of the species of Ahaetulla from the Indo-Malayan Region is proposed. This addition brings to 134 the number of snake species currently known from Sumatra Island.

Phylogenetic relationships among the species of Mabuya present around the Caribbean Sea (Antilles, Central America, and northern South America) are proposed for the first time. The molecular phylogenetic analyses (based on cytochrome b and 12S sequences) give new insights into the diversification of the genus in the New World, its multiple instances of montane habitat colonization, and the multiple waves of colonization towards the Caribbean area. In addition to the molecular analyses, we propose hypotheses about the phylogenetic placement of some rare or possibly extinct species, based on a qualitative analysis of morphological characters. The present article also includes descriptions of two new species from northern Venezuela, Mabuya nebulosylvestrissp. nov. (from the highlands of the coastal range and the Andean Cordillera of Merida) and Mabuya zuliae sp. nov. (from the lowlands of the Maracaibo lake basin). Mabuya luciaeGarman, 1887, a possibly extinct species endemic to St Lucia Island, is resurrected and diagnosed from its supposed sister species, Mabuya mabouya (Lacepede, 1788). © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 598–616.

Three enigmatic skinks, Mabuya nigropalmata Andersson, 1918, Tiliqua maculata Gray, 1839, and Trachylepis (Xystrolepis) punctata, Tschudi, 1845, have been reported from widely separated localities on the South American mainland. They remain the three rarest and most poorly known South American skinks and were known only from type material until recently. In this paper, we comment on recent rediscoveries of M. nigropalmata in the western Amazon of Peru and Bolivia. Using cytochrome b and 12S DNA sequences, we resolve this species’s phylogenetic position within Mabuya sensu stricto and demonstrate its specific distinctiveness, especially from the superficially similar species M. frenata. In addition, we show that neither Trachylepis (Xystrolepis) punctata nor Tiliqua maculata can be placed within Mabuya. A suite of morphological characteristics requires the transfer of these two species to the mostly Afro-Malagasy genus Trachylepis. To correct the secondary homonymy of Tschudi’s name, we propose a replacement name for T. punctata. We designate NRM 23258A as the lectotype of Mabuya nigropalmata Andersson and MNHN 2932 as the lectotype of Trachylepis maculata (Gray) new combination. Finally, we propose an identification key to Mabuya of the Amazon basin and adjacent regions.

We provide an updated molecular phylogenetic analysis of global diversity of typhlopid and xenotyphlopid blindsnakes, adding a set of Madagascan samples and sequences of an additional mitochondrial gene to an existing supermatrix of nuclear and mitochondrial gene segments. Our data suggest monophyly of Madagascan typhlopids, exclusive of introduced Indotyphlops braminus. The Madagascar-endemic typhlopid clade includes two species previously assigned to the genus Lemuriatyphlops (in the subfamily Asiatyphlopinae), which were not each others closest relatives. This contradicts a previous study that described Lemuriatyphlops based on a sequence of the cytochrome oxidase subunit 1 gene from a single species and found this species not forming a clade with the other Malagasy species included. Based on our novel phylogenetic assessment we include all species in this endemic typhlopid clade in the genus Madatyphlops and in the subfamily Madatyphlopinae and consider Lemuriatyphlops as junior synonym. Within Madatyphlops, we identify several candidate species. For some of these (those in the M. arenarius complex), our preliminary data suggest sympatric occurrence and morphological differentiation, thus the existence of undescribed species. We also comment on the genus-level classification of several non-Madagascan typhlopids. We suggest that African species included in Madatyphlops (Afrotyphlops calabresii, A. cuneirostris, A. platyrhynchus, and Rhinotyphlops leucocephalus) should not be included in this genus. We furthermore argue that recent claims of Sundatyphlops, Antillotyphlops, and Cubatyphlops being "undiagnosable" or "not monophyletic" were based on errors in tree reconstruction and failure to notice diagnostic characters, and thus regard these three genera as valid.

We describe a new leaf-tailed gecko species of the Uroplatus ebenaui group from the eastern central rainforests of Madagascar, which had previously been considered as a confirmed candidate species. Our description of Uroplatus fiera sp. nov. relies on integrating evidence from molecular and morphological characters and is based on newly collected material from two localities. A phylogenetic analysis based on multiple mitochondrial DNA fragments places the new species as sister to a lineage of uncertain status (Uroplatus ebenaui [Ca8]), and the clade consisting of these two lineages is sister to a further undescribed candidate species (U. ebenaui [Ca1]). This entire clade is sister to U. phantasticus plus another candidate species. The new species differs from these close relatives, and all other congenerics, by strong differences in DNA sequences of mitochondrial genes (gt;8.5% uncorrected p-distance in 16S rDNA to all nominal species of the genus) and lacks shared alleles with any of the nominal species in the nuclear CMOS gene. From its closest relatives the new species further differs in its much smaller tail size (relative to U. phantasticus), and a narrower tail, fewer supralabials, and more toe lamellae (relative to U. ebenaui [Ca1]). Morphologically the new species is most similar to U. ebenaui but differs in its larger body size and unpigmented oral mucosa. Given its distribution in central eastern Madagascar, with records from near Fierenana and Ambatovy, its range overlaps with that of U. phantasticus. Based on examination of the U. phantasticus holotype, we confirm that this latter has a blackish pigmented oral mucosa as do those specimens typically attributed to this nomen, thereby confirming its distinctness from U. fiera sp. nov., in which the mucosa is unpigmented.

Understanding the phylogenetic and geographical history of Neotropical lineages requires having adequate geographic and taxonomic sampling across the region. However, Colombia has remained a geographical gap in many studies of Neotropical diversity. Here we present a study of Neotropical skinks of the genus Mabuya, reptiles that are difficult to identify or delimit due to their conservative morphology. The goal of the present study is to propose phylogenetic and biogeographic hypotheses of Mabuya including samples from the previously under-studied territory of Colombia, and address relevant biogeographic and taxonomic issues. We combined molecular and morphological data sampled densely by us within Colombia with published data representing broad sampling across the Neotropical realm, including DNA sequence data from two mitochondrial (12S rRNA and cytochrome b) and three nuclear genes (Rag2, NGFB and R35). To evaluate species boundaries we employed a general mixed Yule-coalescent (GMYC) model applied to the mitochondrial data set. Our results suggest that the diversity of Mabuya within Colombia is higher than previously recognized, and includes lineages from Central America and from eastern and southern South America. The genus appears to have originated in eastern South America in the Early Miocene, with subsequent expansions into Central America and the Caribbean in the Late Miocene, including at least six oceanic dispersal events to Caribbean Islands. We identified at least four new candidate species for Colombia and two species that were not previously reported in Colombia. The populations of northeastern Colombia can be assigned to M. zuliae, while specimens from Orinoquia and the eastern foothills of the Cordillera Oriental of Colombia correspond to M. altamazonica. The validity of seven species of Mabuya sensu lato was not supported due to a combination of three factors: (1) non-monophyly, (2)

Scincine lizards in Madagascar form an endemic clade of about 60 species exhibiting a variety of ecomorphological adaptations. Several subclades have adapted to burrowing and convergently regressed their limbs and eyes, resulting in a variety of partial and completely limbless morphologies among extant taxa. However, patterns of limb regression in these taxa have not been studied in detail. Here we fill this gap in knowledge by providing a phylogenetic analysis of DNA sequences of three mitochondrial and four nuclear gene fragments in an extended sampling of Malagasy skinks, and microtomographic analyses of osteology of various burrowing taxa adapted to sand substrate. Based on our data we propose to (i) consider Sirenoscincus Sakata & Hikida, 2003, as junior synonym of Voeltzkowia Boettger, 1893; (ii) resurrect the genus name Grandidierina Mocquard, 1894, for four species previously included in Voeltzkowia; and (iii) consider Androngo Brygoo, 1982, as junior synonym of Pygomeles Grandidier, 1867. By supporting the clade consisting of the limbless Voeltzkowia mira and the forelimb-only taxa V. mobydick and V. yamagishii, our data indicate that full regression of limbs and eyes occurred in parallel twice in the genus Voeltzkowia (as hitherto defined) that we consider as a sand-swimming ecomorph: in the Voeltzkowia clade sensu stricto the regression first affected the hindlimbs and subsequently the forelimbs, whereas the Grandidierina clade first regressed the forelimbs and subsequently the hindlimbs following the pattern prevalent in squamates. Timetree reconstructions for the Malagasy Scincidae contain a substantial amount of uncertainty due to the absence of suitable primary fossil calibrations. However, our preliminary reconstructions suggest rapid limb regression in Malagasy scincids with an estimated maximal duration of 6 MYr for a complete regression in Paracontias, and 4 and 8 MYr respectively for complete regression of forelimbs in Grandidierina and hindlimbs in Voeltzkowia.

The sporadic occurrence of localised pits on parietal plates was recently discovered in different colubrid genera of the subfamily Psammophiinae; these were considered to play a role in sensory perception. In the present study, we describe the presence of similar structures in Atretium schistosum , another colubrid snake reportedly not belonging to the Psammophiinae. As this species is suspected of being phylogenetically distantly related to psammophine snakes, some hypotheses are provided to explain (1) the putative function of these pits, (2) their sporadic occurrence, and (3) to suggest when they may have evolved in the colubroid snake clade. To cite this article: A. Miralles, I. Ineich, C. R. Biologies 329 (2006).

Delimiting and describing species is fundamental to numerous biological disciplines such as evolution, macroecology, and conservation. Delimiting species as independent evolutionary lineages may and often does yield different outcomes depending on the species criteria applied, but methods should be chosen that minimize the inference of objectively erroneous species limits. Several protocols exploit single-gene or multi-gene coalescence statistics, assignment tests or other rationales related to nuclear DNA (nDNA) allele sharing to automatically delimit species. We apply seven different species delimitation protocols to a taxonomically confusing group of Malagasy lizards (Madascincus), and compare the resulting taxonomies with two newly developed metrics: the Taxonomic index of congruence C tax which quantifies the congruence between two taxonomies, and the Relative taxonomic resolving power index R tax which quantifies the potential of an approach to capture a high number of species boundaries. The protocols differed in the total number of species proposed, between 9 and 34, and were also highly incongruent in placing species boundaries. The Generalized Mixed Yule-Coalescent approach captured the highest number of potential species boundaries but many of these were clearly contradicted by extensive nDNA admixture between sympatric mitochondrial DNA (mtDNA) haplotype lineages. Delimiting species as phenotypically diagnosable mtDNA clades failed to detect two cryptic species that are unambiguous due to a lack of nDNA gene flow despite sympatry. We also consider the high number of species boundaries and their placement by multi-gene Bayesian species delimitation as poorly reliable whereas the Bayesian assignment test approach provided a species delimitation highly congruent with integrative taxonomic practice. The present study illustrates the trade-off in taxonomy between reliability (favored by conservative approaches) and resolving power (favored by inflationist approaches). Quantifying excessive splitting is more difficult than quantifying excessive lumping, suggesting a priority for conservative taxonomies in which errors are more liable to be detected and corrected by subsequent studies.

The Linnaean classification system provides the universal reference system for communicating about the diversity of life and its hierarchic history. Several limitations that challenge the stability of this system have been identified and, as a result, alternative systems have been proposed since its early inception. The revolution caused by molecular phylogenetics has, more than ever, exemplified that Linnaean classification schemes are subject to a degree of instability that may hamper their significance and communication power. Our analysis of recent changes in the classification of several groups of organisms, with a focus on amphibians and reptiles, reveals two main sources of instability: (i) revisionary, objective (empirical) changes based on the discovery of unambiguous instances of non-monophyly and on progress in the Globe’s species inventory, and (ii) subjective changes based on author preferences or on a poor analysis of the advantages and limitations of new classification schemes. To avoid subjective taxonomic instability, we review and elaborate proposals for the assignment of Linnaean rank to clades, and thereby for the naming of these clades as Linnaean taxa (Taxon Naming Criteria: TNCs). These are drafted from the perspective of practicing taxonomists and can help choosing among alternative monophyly-based classifications under a premise of economy of change. We provide a rationale for each TNC along with real and theoretical examples to illustrate their practical advantages and disadvantages. We conclude that not all TNCs lead to equally informative and stable taxonomies. Therefore, we order the various TNCs by the generality of their implications and provide a workflow scheme to guide the procedure of taxonomic decisions concerning the creation or modification of supraspecific classifications. The following criteria are considered primary when naming taxa: (i) Mono phyly of the taxon in an inferred species tree; (ii) Clade Stability, i.e., the monophyly of a clade to be named as taxon should be as strongly supported as possible by various methods of tree inference, tests of clade robustness, and different data sets; and (iii) Phenotypic Diagnosability, i.e., ranked supraspecific taxa should be those that are phenotypically most conspicuous although in phenotypically cryptic groups of organisms it can be warranted to name taxa based on molecular differences alone. We consider various other criteria as secondary (i.e., the Time Banding, Biogeography, Adaptive Zone, and Hybrid Viability TNCs) and refute using them as sole arguments for the modification of established classifications or proposal of new ones. Taxonomists are encouraged to be explicit and consistent when applying TNCs for creating or modifying classifications. We emphasize that, except for monophyly, the priority TNCs are not proposed as mandatory requisites of a Linnaean taxon but as yardsticks to allow for an informed choice among various clades in a tree that could alternatively be named as Linnaean taxa. Despite a need for plurality, classifications should avoid deliberately violating any of the three primary TNCs because taxa of unstable monophyly or poor diagnosability reduce the information content and hence the utility of the Linnaean system.