The Museo Civico di Storia Naturale ‘‘Giacomo Doria,’’ Genoa, hosts one of the most important historical collections of the Opiliones subor- der Cyphophthalmi, including all the known specimens for the type species of the genera Leptopsalis (Stylo- cellidae), Miopsalis (Stylocellidae), and Parogovia (Neo- goveidae), as well as several other types in the families Ogoveidae and Stylocellidae (it is unclear whether specimens in Pettalidae and Sironidae constitute types). These specimens were recently made available to us for study, and given their importance, we discuss and illustrate them here. Study of this collection allows confirmation of the validity of Leptopsalis, considered a synonym of Stylocellus for more than a century, and of Miopsalis, considered a nomen dubium in the most recent catalogue of the group. It furthermore helps to clarify the identity of several other species in the family Stylocellidae. Here we formally resurrect the genera Miopsalis Thorell, 1890, and Leptopsalis Thorell, 1882, and transfer several species to these genera: M. collinsi (Shear, 1993) comb. nov.; M. gryllospeca (Shear, 1993) comb. nov.; M. lionota (Pocock, 1897) comb. nov.; M. sabah (Shear, 1993) comb. nov.; M. silhavyi (Rambla, 1991) comb. nov.; M. tarumpitao (Shear, 1993) comb. nov.; L. dumoga (Shear, 1993) comb. nov.; L. hillyardi (Shear, 1993) comb. nov.; L. javana Thorell, 1882; L. lydekkeri (Clouse & Giribet, 2007) comb. nov.; L. modesta (Hansen & Sørensen, 1904) comb. nov.; L. novaguinea (Clouse & Giribet, 2007) comb. nov.; L. ramblae (Giribet, 2002) comb. nov.; L. sulcata (Hansen & Sørensen, 1904) comb. nov.; L. tambusisi (Shear, 1993) comb. nov.; L. thorellii (Hansen & Sørensen, 1904) comb. nov.; L. weberii (Hansen & Sørensen, 1904) comb. nov.

The phylogenetic relationships of selected members of the phylum Nemertea are explored by means of six markers amplified from the genomic DNA of freshly collected specimens (the nuclear 18S rRNA and 28S rRNA genes, histones H3 and H4, and the mitochondrial genes 16S rRNA and cytochrome c oxidase subunit I). These include all previous markers and regions used in earlier phylogenetic analyses of nemerteans, therefore acting as a scaffold to which one could pinpoint any previously published study. Our results, based on analyses of static and dynamic homology concepts under probabilistic and parsimony frameworks, agree in the non-monophyly of Palaeonemertea and in the monophyly of Heteronemerta and Hoplonemertea. The position of Hubrechtella and the Pilidiophora hypothesis are, however, sensitive to analytical method, as is the monophyly of the non-hubrechtiid palaeonemerteans. Our results are, however, consistent with the main division of Hoplonemertea into Polystilifera and Monostilifera, the last named being divided into Cratenemertea and Distromatonemertea, as well as into the main division of Heteronemertea into Baseodiscus and the remaining species. The study also continues to highlight the deficient taxonomy at the family and generic level within Nemertea and sheds light on the areas of the tree that require further refinement.

Identifying the unambiguous members of Vetigastropoda and understanding the relationships among its families has been challenging. This study investigates the internal relationships among putative members of Vetigastropoda sensu lato (Fissurelloidea, Haliotoidea, Lepetelloidea, Lepetodriloidea, Pleurotomarioidea, Scissurelloidea, Seguenzioidea, Trochoidea, Angarioidea, Phasianelloidea, Neomphaloidea and Cocculinoidea) in a molecular phylogeny utilizing nearly 6 kb of molecular data from up to five nuclear and mitochondrial genes. Single-step parsimony-based and two-step maximum-likelihood analyses are employed as phylogenetic methods to analyse the data. Sequence data from all vetigastropod groups are included and in order to overcome shortfalls of previous vetigas- tropod analyses resulting from the under-sampling of outgroups, this study also includes broad out- group representation. Fissurelloidea, Haliotoidea, Lepetodriloidea, Scissurelloidea, Seguenzioidea, Trochoidea, Angarioidea and Phasianelloidea formed a clade identified as Vetigastropoda sensu stricto united by morphological synapomorphies such as the presence of bursicles and epipodial sense organs. In contrast, Neomphalina, Cocculinoidea and Pleurotomarioidea fell outside Vetigastropoda s. s., indi- cating a need to reexamine the classification of these clades as vetigastropods.

We investigate the phylogeny, biogeography, time of origin and diversification, ancestral area reconstruction and large-scale distributional patterns of an ancient group of arachnids, the harvestman suborder Cyphophthalmi. Analysis of molecular and morphological data allow us to propose a new classification system for the group; Pettalidae constitutes the infraorder Scopulophthalmi new clade, sister group to all other families, which are divided into the infraorders Sternophthalmi new clade and Boreophthalmi new clade. Sternophthalmi includes the families Troglosironidae, Ogoveidae, and Neogoveidae; Boreophthalmi includes Stylocellidae and Sironidae, the latter family of questionable monophyly. The internal resolution of each family is discussed and traced back to its geological time origin, as well as to its original landmass, using methods for estimating divergence times and ancestral area reconstruction. The origin of Cyphophthalmi can be traced back to the Carboniferous, whereas the diversification time of most families ranges between the Carboniferous and the Jurassic, with the exception of Troglosironidae, whose current diversity originates in the Cretaceous/Tertiary. Ancestral area reconstruction is ambiguous in most cases. Sternophthalmi is traced back to an ancestral land mass that contained New Caledonia and West Africa in the Permian, whereas the ancestral landmass for Neogoveidae included the south-eastern USA and West Africa, dating back to the Triassic. For Pettalidae, most results include South Africa, or a combination of South Africa with the Australian plate of New Zealand or Sri Lanka, as the most likely ancestral landmass, back in the Jurassic. Stylocellidae is reconstructed to the Thai-Malay Penisula during the Jurassic. Combination of the molecular and morphological data results in a hypothesis for all the cyphophthalmid genera, although the limited data available for some taxa represented only in the morphological partition negatively affects the phylogenetic reconstruction by decreasing nodal support in most clades. However, it resolves the position of many monotypic genera not available for molecular analysis, such as Iberosiro, Odontosiro, Speleosiro, Managotria or Marwe, although it does not place Shearogovea or Ankaratra within any existing family. The biogeographical data show a strong correlation between relatedness and formerly adjacent landmasses, and oceanic dispersal does not need to be postulated to explain disjunct distributions, especially when considering the time of divergence. The data also allow testing of the hypotheses of the supposed total submersion of New Zealand and New Caledonia, clearly falsifying submersion of the former, although the data cannot reject the latter.

Sclerosomatids constitute the largest family of the arachnid order Opiliones, and one of the two families com- monly found in the temperate regions of the northern Hemi- sphere. Harvestmen have a sparse fossil record in the Mesozoic, with only two species known from the Jurassic, one of them poorly preserved and none with precise phylo- genetic placement. Here we report a new fossil, Mesobunus dunlopi sp. nov., from the Middle Jurassic (approx. 165 Mya) of Daohugou, Inner Mongolia, China. The new spe- cies is related to another genus of the same formation, but the preservation quality and details of the penis and pedi- palps allow us to place them in the extant sclerosomatid subfamilies Gagrellinae or Leiobuninae. The first recognis- able fossil in this subfamily highlights morphological stasis over ca. 165 Mya and the finding of this species along with lacustrine insects suggests a life mode similar to that of some modern sclerosomatids, and a possible connection between morphological and ecological stasis.

Coral reefs are increasingly threatened worldwide by a variety of biological and physical factors, including disease, bleaching and ocean acidification. Under- standing levels of connectivity among widespread populations can assist in con- servation efforts and the design of marine protected areas, as larval dispersal scales affect population demography. This study examined genetic connectivity and morphological variation of the broadcast spawning coral Montastraea cavernosa (L., 1767) among five locations in the Caribbean and Western Atlan- tic. Analysis of mtDNA and nuclear rRNA internal transcribed spacers, at both the local and regional scale, revealed that the majority of variation existed within locations rather than among them. Likewise, the majority of pairwise compari- sons were non-significant between sites and locations. These results suggest that moderate to high gene flow occurs within and among populations of M. cavern- osa in the Western Atlantic. The phylogeographic signature and significant pair- wise comparisons among several locations, however, indicate that populations are also partially maintained through self-seeding and that gene flow may be restricted over large geographic distances. Additionally, while some anatomical variation is likely attributable to phenotypic plasticity, variations in skeletal mor- phology between Jamaica and other locations correspond with significant pair- wise genetic distances and the presence of private sequence types (limited to a single location), suggesting selection to local environmental conditions.

Arthropods are the most diverse group of animals and have been so sincethe Cambrian radiation. They belong to the protostome clade Ecdysozoa,with Onychophora (velvet worms) as their most likely sister group andtardigrades (water bears) the next closest relative. The arthropod tree oflife can be interpreted as a five-taxon network, containing Pycnogonida,Euchelicerata, Myriapoda, Crustacea, and Hexapoda, the last two formingthe clade Tetraconata or Pancrustacea. The unrooted relationship of Tetraconatato the three other lineages is well established, but of three possiblerooting positions the Mandibulata hypothesis receives the most support.Novel approaches to studying anatomy with noninvasive three-dimensionalreconstruction techniques, the application of these techniques to new andold fossils, and the so-called next-generation sequencing techniques are atthe forefront of understanding arthropod relationships. Cambrian fossilsassigned to the arthropod stem group inform on the origin of arthropodcharacters from a lobopodian ancestry. Monophyly of Pycnogonida, Euchelicerata,Myriapoda, Tetraconata, and Hexapoda is well supported, but theinterrelationships of arachnid orders and the details of crustacean paraphylywith respect to Hexapoda remain themajor unsolved phylogenetic problems.

The first scolopocryptopid centipede known from the fossil record is a specimen of the subfamily Scolopocryp- topinae in Miocene amber from Chiapas, southern Mexico. It is described here as Scolopocryptops simojo- velensis sp. nov., displaying a distinct combination of morphological characters compared to extant congeners. Anatomical details of the fossil specimen were acquired by non-invasive 3D synchrotron microtomography using X-ray phase contrast. The phylogenetic position of the new species is inferred based on a combination of morphological data with sequences for six genes (nuclear 18S and 28S rRNA, nuclear protein-coding histone H3, and mitochondrial 12S rRNA, 16S rRNA, and protein-coding cytochrome c oxidase subunit I) for extant Scolopendromorpha. The data set includes eight extant species of Scolopocryptops and Dinocryptops from North America, east Asia, and the Pacific, rooted with novel sequence data for other blind scolopendromorphs. The molecular and combined data sets, analysed in a parsimony/direct optimization framework, identified a stable pattern of two main clades within Scolopocryptopinae. North American and Asian species of Scolopoc- ryptops are united as a clade supported by both morphological and molecular characters. Its sister group is a Neotropical clade in which the type species of Dinocryptops is nested within a paraphyletic assemblage of Scolopocryptops species; Dinocryptops is placed in synonymy with Scolopocryptops. The strength of support for the relationships of extant taxa from the molecular data allow the Chiapas fossil to be assigned with precision, despite ambiguity in the morphological data; the fossil is resolved as sister species to the extant Laurasian clade.

In order to study the tempo and the mode of spider orb web evolution and diversification, we conducted a phylogenetic analysis using six genetic markers along with a comprehensive taxon sample. The present analyses are the first to recover the monophyly of orb-weaving spiders based solely on DNA sequence data and an extensive taxon sample. We present the first dated orb weaver phylogeny. Our results suggest that orb weavers appeared by the Middle Triassic and underwent a rapid diversification during the end of the Triassic and Early Jurassic. By the second half of the Jurassic, most of the extant orb-weaving families and web designs were already present. The processes that may have given origin to this diversification of lineages and web architectures are discussed. A combination of biotic factors, such as key innovations in web design and silk composition, as well as abiotic environmental changes, may have played important roles in the diversification of orb weavers. Our analyses also show that increased taxon sampling density in both ingroups and outgroups greatly improves phylogenetic accuracy even when extensive data are missing. This effect is particularly important when addition of character data improves gene overlap.

Many recent studies on invertebrates have shown how morphology not always captures the true diversity of taxa, with cryptic speciation often being discussed in this context. Here, we show how diversification patterns can be very different in two clades of closely related earthworms in the genus Hormogaster stressing the risk of using nonspecific substitution rate values across taxa. On the one hand, the Hormogaster elisae species complex, endemic to the central Iberian Peninsula, shows morphological stasis. On the other hand, a clade of Hormogaster from the NE Iberian Peninsula shows an enormous morphological variability, with 15 described morphospecies. The H. elisae complex, however, evolves faster genetically, and this could be explained by the harsher environmental conditions to which it is confined—as detected in this study, that is, sandier and slightly poorer soils with lower pH values than those of the other species in the family. These extreme conditions could be at the same time limiting morphological evolution and thus be responsible for the observed morphological stasis in this clade. Contrarily, Hormogaster species from the NE Iberian Peninsula, although still inhabiting harsher milieu than other earthworm groups, have had the opportunity to evolve into a greater morphological disparity. An attempt to delimit species within this group following the recently proposed general mixed Yule-coalescent method showed a higher number of entities than expected under the morphospecies concept, most probably due to the low vagility of these animals, which considerably limits gene flow between distant conspecific populations, but also because of the decoupling between morphological and genetic evolution in the H. elisae complex.

Tardigrades constitute a phylum of miniaturized metazoans with ca. 1030 living species, a fossil record that probably dates back to the Cambrian, and physiological properties that allow them to live in almost any environment known to host life on Earth—they can also survive in space. Despite broad consensus regarding their membership of the superclade Ecdysozoa, the exact position of the phylum remains contested (some analyses suggest onychophorans and arthropods as their closest relatives, while others suggest a relationship to nematodes and nematomorphs) and the internal relationships of the phylum are still poorly understood. In the present study, we present a hypothesis of tardigrade relationships by examining more taxa and more markers than any previously published phylogeny of the group. We generated novel data for three markers (18S rRNA, 28S rRNA, COI) for 42 individuals of 16 carefully identified species, comprising 12 genera and five families from the classes Heterotardigrada and Eutardigrada, and analysed them in conjunction with nearly all data available from GenBank. Our results show certain disagreement with current taxonomy both at higher ranks (families, orders, classes) and at low (generic) taxonomic levels. When studying the sensitivity to outgroup choice, the class Eutardigrada was monophyletic under only one combination of outgroups; all other combinations placed the eutardigrade order Apochela as sister to the class Heterotardigrada. Phylogenetic relationships within the other eutardigrade order, Parachela, were stable to outgroup choice. Eutardigrade superfamilies recently proposed by Sands and collaborators in the order Parachela were tested with the introduction of new sequences from additional genera, and the possible morphological synapomorphies supporting those superfamilies are discussed.