Molecular systematics and phylogenetics of the spider genus Mastigusa Menge, 1854 (Araneae, Cybaeidae).

The palearctic spider genus Mastigusa Menge, 1854 is characterized by a remarkable morphology and wide ecological variability, with free-living, cave dwelling and myrmecophile populations known. This genus has a long and tangled taxonomic history and was placed in different families in the past, all belonging to the "marronoid clade", an informal grouping of families characterized by the lack of strong synapomorphies. Three species are currently recognized, but their identity and circumscription has been long debated. A molecular approach was never applied for trying to solve these uncertainties, and doubts still remain both about its phylogenetic placement and about the taxonomic status of the described species. For the first time the genus Mastigusa is included in a molecular phylogenetic analysis and strong support is found for its placement within the family Cybaeidae, in sister relationship with the genus Cryphoeca Thorell, 1870. An analysis of Mastigusa populations spanning across the distribution range of the genus identifies a high and previously overlooked genetic diversity, with six distinct genetic lineages showing a strong geographic pattern. Divergence times between Mastigusa and its sister genus and between the distinct Mastigusa lineages are estimated, and the groundwork is laid for a taxonomic revision of the species belonging to the genus.

Phylogeny of the orb-weaving spider family Araneidae (Araneae: Araneoidea).

We present a new phylogeny of the spider family Araneidae based on five genes (28S, 18S, COI, H3 and 16S) for 158 taxa, identified and mainly sequenced by us. This includes 25 outgroups and 133 araneid ingroups representing the subfamilies Zygiellinae Simon, 1929, Nephilinae Simon, 1894, and the typical araneids, here informally named the "ARA Clade". The araneid genera analysed here include roughly 90% of all currently named araneid species. The ARA Clade is the primary focus of this analysis. In taxonomic terms, outgroups comprise 22 genera and 11 families, and the ingroup comprises three Zygiellinae and four Nephilinae genera, and 85 ARA Clade genera (ten new). Within the ARA Clade, we recognize ten informal groups that contain at least three genera each and are supported under Bayesian posterior probabilities (≥ 0.95): "Caerostrines" (Caerostris, Gnolus and Testudinaria), "Micrathenines" (Acacesia, Micrathena, Ocrepeira, Scoloderus and Verrucosa), "Eriophorines" (Acanthepeira, Alpaida, Eriophora, Parawixia and Wagneriana), "Backobourkiines" (Acroaspis, Backobourkia, Carepalxis, Novakiella, Parawixia, Plebs, Singa and three new genera), "Argiopines" (Arachnura, Acusilas, Argiope, Cyrtophora, Gea, Lariniaria and Mecynogea), "Cyrtarachnines" (Aranoethra, Cyrtarachne, Paraplectana, Pasilobus and Poecilopachys), "Mastophorines" (Celaenia, Exechocentrus and Mastophora,), "Nuctenines" (Larinia, Larinioides and Nuctenea), "Zealaraneines" (Colaranea, Cryptaranea, Paralarinia, Zealaranea and two new genera) and "Gasteracanthines" (Augusta, Acrosomoides, Austracantha, Gasteracantha, Isoxya, Macracantha, Madacantha, Parmatergus and Thelacantha). Few of these groups are currently corroborated by morphology, behaviour, natural history or biogeography. We also include the large genus Araneus, along with Aculepeira, Agalenatea, Anepsion, Araniella, Cercidia, Chorizopes, Cyclosa, Dolophones, Eriovixia, Eustala, Gibbaranea, Hingstepeira, Hypognatha, Kaira, Larinia, Mangora, Metazygia, Metepeira, Neoscona, Paraplectanoides, Perilla, Poltys, Pycnacantha, Spilasma and Telaprocera, but the placement of these genera was generally ambiguous, except for Paraplectanoides, which is strongly supported as sister to traditional Nephilinae. Araneus, Argiope, Eriophora and Larinia are polyphyletic, Araneus implying nine new taxa of genus rank, and Eriophora and Larinia two each. In Araneus and Eriophora, polyphyly was usually due to north temperate generic concepts being used as dumping grounds for species from southern hemisphere regions, e.g. South-East Asia, Australia or New Zealand. Although Araneidae is one of the better studied spider families, too little natural history and/or morphological data are available across these terminals to draw any strong evolutionary conclusions. However, the classical orb web is reconstructed as plesiomorphic for Araneidae, with a single loss in "cyrtarachnines"-"mastophorines". Web decorations (collectively known as stabilimenta) evolved perhaps five times. Sexual dimorphism generally results from female body size increase with few exceptions; dimorphic taxa are not monophyletic and revert to monomorphism in a few cases.

Next-generation museum genomics: Phylogenetic relationships among palpimanoid spiders using sequence capture techniques (Araneae: Palpimanoidea).

Historical museum specimens are invaluable for morphological and taxonomic research, but typically the DNA is degraded making traditional sequencing techniques difficult to impossible for many specimens. Recent advances in Next-Generation Sequencing, specifically target capture, makes use of short fragment sizes typical of degraded DNA, opening up the possibilities for gathering genomic data from museum specimens. This study uses museum specimens and recent target capture sequencing techniques to sequence both Ultra-Conserved Elements (UCE) and exonic regions for lineages that span the modern spiders, Araneomorphae, with a focus on Palpimanoidea. While many previous studies have used target capture techniques on dried museum specimens (for example, skins, pinned insects), this study includes specimens that were collected over the last two decades and stored in 70% ethanol at room temperature. Our findings support the utility of target capture methods for examining deep relationships within Araneomorphae: sequences from both UCE and exonic loci were important for resolving relationships; a monophyletic Palpimanoidea was recovered in many analyses and there was strong support for family and generic-level palpimanoid relationships. Ancestral character state reconstructions reveal that the highly modified carapace observed in mecysmaucheniids and archaeids has evolved independently.

Rounding up the usual suspects: a standard target-gene approach for resolving the interfamilial phylogenetic relationships of ecribellate orb-weaving spiders with a new family-rank classification (Araneae, Araneoidea).

We test the limits of the spider superfamily Araneoidea and reconstruct its interfamilial relationships using standard molecular markers. The taxon sample (363 terminals) comprises for the first time representatives of all araneoid families, including the first molecular data of the family Synaphridae. We use the resulting phylogenetic framework to study web evolution in araneoids. Araneoidea is monophyletic and sister to Nicodamoidea rank. n. Orbiculariae are not monophyletic and also include the RTA clade, Oecobiidae and Hersiliidae. Deinopoidea is paraphyletic with respect to a lineage that includes the RTA clade, Hersiliidae and Oecobiidae. The cribellate orb-weaving family Uloboridae is monophyletic and is sister group to a lineage that includes the RTA Clade, Hersiliidae and Oecobiidae. The monophyly of most Araneoidea families is well supported, with a few exceptions. Anapidae includes holarchaeids but the family remains diphyletic even if Holarchaea is considered an anapid. The orb-web is ancient, having evolved by the early Jurassic; a single origin of the orb with multiple "losses" is implied by our analyses. By the late Jurassic, the orb-web had already been transformed into different architectures, but the ancestors of the RTA clade probably built orb-webs. We also find further support for a single origin of the cribellum and multiple independent losses. The following taxonomic and nomenclatural changes are proposed: the cribellate and ecribellate nicodamids are grouped in the superfamily Nicodamoidea rank n. (Megadictynidae rank res. and Nicodamidae stat. n.). Araneoidea includes 17 families with the following changes: Araneidae is re-circumscribed to include nephilines, Nephilinae rank res., Arkyidae rank n., Physoglenidae rank n., Synotaxidae is limited to the genus Synotaxus, Pararchaeidae is a junior synonym of Malkaridae (syn. n.), Holarchaeidae of Anapidae (syn. n.) and Sinopimoidae of Linyphiidae (syn. n.).

The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling.

We present a phylogenetic analysis of spiders using a dataset of 932 spider species, representing 115 families (only the family Synaphridae is unrepresented), 700 known genera, and additional representatives of 26 unidentified or undescribed genera. Eleven genera of the orders Amblypygi, Palpigradi, Schizomida and Uropygi are included as outgroups. The dataset includes six markers from the mitochondrial (12S, 16S, COI) and nuclear (histone H3, 18S, 28S) genomes, and was analysed by multiple methods, including constrained analyses using a highly supported backbone tree from transcriptomic data. We recover most of the higher-level structure of the spider tree with good support, including Mesothelae, Opisthothelae, Mygalomorphae and Araneomorphae. Several of our analyses recover Hypochilidae and Filistatidae as sister groups, as suggested by previous transcriptomic analyses. The Synspermiata are robustly supported, and the families Trogloraptoridae and Caponiidae are found as sister to the Dysderoidea. Our results support the Lost Tracheae clade, including Pholcidae, Tetrablemmidae, Diguetidae, Plectreuridae and the family Pacullidae (restored status) separate from Tetrablemmidae. The Scytodoidea include Ochyroceratidae along with Sicariidae, Scytodidae, Drymusidae and Periegopidae; our results are inconclusive about the separation of these last two families. We did not recover monophyletic Austrochiloidea and Leptonetidae, but our data suggest that both groups are more closely related to the Cylindrical Gland Spigot clade rather than to Synspermiata. Palpimanoidea is not recovered by our analyses, but also not strongly contradicted. We find support for Entelegynae and Oecobioidea (Oecobiidae plus Hersiliidae), and ambiguous placement of cribellate orb-weavers, compatible with their non-monophyly. Nicodamoidea (Nicodamidae plus Megadictynidae) and Araneoidea composition and relationships are consistent with recent analyses. We did not obtain resolution for the titanoecoids (Titanoecidae and Phyxelididae), but the Retrolateral Tibial Apophysis clade is well supported. Penestomidae, and probably Homalonychidae, are part of Zodarioidea, although the latter family was set apart by recent transcriptomic analyses. Our data support a large group that we call the marronoid clade (including the families Amaurobiidae, Desidae, Dictynidae, Hahniidae, Stiphidiidae, Agelenidae and Toxopidae). The circumscription of most marronoid families is redefined here. Amaurobiidae include the Amaurobiinae and provisionally Macrobuninae. We transfer Malenellinae (Malenella, from Anyphaenidae), Chummidae (Chumma) (new syn.) and Tasmarubriinae (Tasmarubrius, Tasmabrochus and Teeatta, from Amphinectidae) to Macrobuninae. Cybaeidae are redefined to include Calymmaria, Cryphoeca, Ethobuella and Willisius (transferred from Hahniidae), and Blabomma and Yorima (transferred from Dictynidae). Cycloctenidae are redefined to include Orepukia (transferred from Agelenidae) and Pakeha and Paravoca (transferred from Amaurobiidae). Desidae are redefined to include five subfamilies: Amphinectinae, with Amphinecta, Mamoea, Maniho, Paramamoea and Rangitata (transferred from Amphinectidae); Ischaleinae, with Bakala and Manjala (transferred from Amaurobiidae) and Ischalea (transferred from Stiphidiidae); Metaltellinae, with Austmusia, Buyina, Calacadia, Cunnawarra, Jalkaraburra, Keera, Magua, Metaltella, Penaoola and Quemusia; Porteriinae (new rank), with Baiami, Cambridgea, Corasoides and Nanocambridgea (transferred from Stiphidiidae); and Desinae, with Desis, and provisionally Poaka (transferred from Amaurobiidae) and Barahna (transferred from Stiphidiidae). Argyroneta is transferred from Cybaeidae to Dictynidae. Cicurina is transferred from Dictynidae to Hahniidae. The genera Neoramia (from Agelenidae) and Aorangia, Marplesia and Neolana (from Amphinectidae) are transferred to Stiphidiidae. The family Toxopidae (restored status) includes two subfamilies: Myroinae, with Gasparia, Gohia, Hulua, Neomyro, Myro, Ommatauxesis and Otagoa (transferred from Desidae); and Toxopinae, with Midgee and Jamara, formerly Midgeeinae, new syn. (transferred from Amaurobiidae) and Hapona, Laestrygones, Lamina, Toxops and Toxopsoides (transferred from Desidae). We obtain a monophyletic Oval Calamistrum clade and Dionycha; Sparassidae, however, are not dionychans, but probably the sister group of those two clades. The composition of the Oval Calamistrum clade is confirmed (including Zoropsidae, Udubidae, Ctenidae, Oxyopidae, Senoculidae, Pisauridae, Trechaleidae, Lycosidae, Psechridae and Thomisidae), affirming previous findings on the uncertain relationships of the "ctenids" Ancylometes and Cupiennius, although a core group of Ctenidae are well supported. Our data were ambiguous as to the monophyly of Oxyopidae. In Dionycha, we found a first split of core Prodidomidae, excluding the Australian Molycriinae, which fall distantly from core prodidomids, among gnaphosoids. The rest of the dionychans form two main groups, Dionycha part A and part B. The former includes much of the Oblique Median Tapetum clade (Trochanteriidae, Gnaphosidae, Gallieniellidae, Phrurolithidae, Trachelidae, Gnaphosidae, Ammoxenidae, Lamponidae and the Molycriinae), and also Anyphaenidae and Clubionidae. Orthobula is transferred from Phrurolithidae to Trachelidae. Our data did not allow for complete resolution for the gnaphosoid families. Dionycha part B includes the families Salticidae, Eutichuridae, Miturgidae, Philodromidae, Viridasiidae, Selenopidae, Corinnidae and Xenoctenidae (new fam., including Xenoctenus, Paravulsor and Odo, transferred from Miturgidae, as well as Incasoctenus from Ctenidae). We confirm the inclusion of Zora (formerly Zoridae) within Miturgidae.

Phantoms of Gondwana?-phylogeny of the spider subfamily Mynogleninae (Araneae: Linyphiidae).

This is the first genus-level phylogeny of the subfamily Mynogleninae. It is based on 190 morphological characters scored for 44 taxa: 37 mynoglenine taxa (ingroup) representing 15 of the 17 known genera and seven outgroup taxa representing the subfamilies Stemonyphantinae, Linyphiinae (Linyphiini and Micronetini), and Erigoninae, and a representative of the family Pimoidae, the sister-group to Linyphiidae. No fewer than 147 of the morphological characters used in this study are new and defined for this study, and come mainly from male and female genitalia. Parsimony analysis with equal weights resulted in three most parsimonious trees of length 871. The monophyly of the subfamily Mynogleninae and the genera Novafroneta, Parafroneta, Laminafroneta, Afroneta, Promynoglenes, Metamynoglenes, and Haplinis are supported, whereas Pseudafroneta is paraphyletic. The remaining seven mynoglenine genera are either monotypic or represented by only one taxon. Diagnoses are given for all genera included in the analysis. The evolution of morphological traits is discussed and we summarize the diversity and distribution patterns of the 124 known species of mynoglenines. The preferred topology suggests a single origin of mynoglenines in New Zealand with two dispersal events to Africa, and does not support Gondwana origin.

The strange case of Laetesia raveni n. sp., a green linyphiid spider from Eastern Australia with a preference for thorny plants (Araneae, Linyphiidae).

Laetesia raveni n. sp. (Araneae, Linyphiidae), is described based on specimens collected in New South Wales and Queensland (Australia). This new linyphiid species is of bright green colour, and it seems to have a preference to build its webs almost exclusively on two plant species, namely Calamus muelleri Wendland (Arecaceae) and Solanum inaequilaterum Domin, (Solanaceae), both of them densely covered with thorns. The epigynal morphology of Laetesia raveni n. sp. varies intraspecifically. Live individuals and several of their dome-shaped sheet webs are illustrated. 

Enigmatic and extravagant genitalia in the spider genus Mastigusa (Araneae, Cybaeidae) - a taxonomic revision.

Mastigusa is a genus of small palearctic spiders that has recently been moved to the family Cybaeidae after the first inclusion of the genus in a phylogenetic matrix. Three species are currently recognised: M. arietina , M. lucifuga and M. macrophthalma . The status and delimitation, though, has always been problematic due to inconsistency in the characters used to discriminate between these, leading to great confusion in identity and distribution. We present a detailed morphological redescription of the genus and a taxonomic revision of the included species by the combined use of morphological data and molecular species-delimitation techniques based on the mitochondrial COI gene. The status of the three currently described species has been reevaluated and Mastigusa diversa was revalidated based on material from the Iberian Peninsula, North Africa and the United Kingdom. The distribution of Mastigusa species is updated based on novel taxonomic considerations, and comments on the natural history and ecological differences observed in the species are provided. ZooBank: urn:lsid:zoobank.org:pub:AAD3FAED-440F-4295-B458-455B1D913F81.

Reconstructing web evolution and spider diversification in the molecular era.

The evolutionary diversification of spiders is attributed to spectacular innovations in silk. Spiders are unique in synthesizing many different kinds of silk, and using silk for a variety of ecological functions throughout their lives, particularly to make prey-catching webs. Here, we construct a broad higher-level phylogeny of spiders combining molecular data with traditional morphological and behavioral characters. We use this phylogeny to test the hypothesis that the spider orb web evolved only once. We then examine spider diversification in relation to different web architectures and silk use. We find strong support for a single origin of orb webs, implying a major shift in the spinning of capture silk and repeated loss or transformation of orb webs. We show that abandonment of costly cribellate capture silk correlates with the 2 major diversification events in spiders (1). Replacement of cribellate silk by aqueous silk glue may explain the greater diversity of modern orb-weaving spiders (Araneoidea) compared with cribellate orb-weaving spiders (Deinopoidea) (2). Within the "RTA clade," which is the sister group to orb-weaving spiders and contains half of all spider diversity, >90% of species richness is associated with repeated loss of cribellate silk and abandonment of prey capture webs. Accompanying cribellum loss in both groups is a release from substrate-constrained webs, whether by aerially suspended webs, or by abandoning webs altogether. These behavioral shifts in silk and web production by spiders thus likely played a key role in the dramatic evolutionary success and ecological dominance of spiders as predators of insects.

Higher-level phylogenetics of linyphiid spiders (Araneae, Linyphiidae) based on morphological and molecular evidence.

This study infers the higher-level cladistic relationships of linyphiid spiders from five genes (mitochondrial CO1, 16S; nuclear 28S, 18S, histone H3) and morphological data. In total, the character matrix includes 47 taxa: 35 linyphiids representing the currently used subfamilies of Linyphiidae (Stemonyphantinae, Mynogleninae, Erigoninae, and Linyphiinae (Micronetini plus Linyphiini)) and 12 outgroup species representing nine araneoid families (Pimoidae, Theridiidae, Nesticidae, Synotaxidae, Cyatholipidae, Mysmenidae, Theridiosomatidae, Tetragnathidae, and Araneidae). The morphological characters include those used in recent studies of linyphiid phylogenetics, covering both genitalic and somatic morphology. Different sequence alignments and analytical methods produce different cladistic hypotheses. Lack of congruence among different analyses is, in part, due to the shifting placement of Labulla, Pityohyphantes, Notholepthyphantes, and Pocobletus. Almost all combined analyses agree on the monophyly of linyphioids, Pimoidae, Linyphiidae, Erigoninae, Mynogleninae, as well as Stemonyphantes as a basal lineage within Linyphiidae. Our results suggest independent origins of the desmitracheate tracheal system in micronetines and erigonines, and that erigonines were primitively haplotracheate. Cephalothoracic glandular specializations of erigonines and mynoglenines apparently evolved independently. Subocular sulci of mynoglenines and lateral sulci (e.g. Bathyphantes) evolved independently but glandular pores in the prosoma proliferated once. The contribution of different character partitions and their sensitivity to changes in traditional analytical parameters is explored and quantified. © The Willi Hennig Society 2009.

A landscape-scale assessment of tropical mammals reveals the effects of habitat and anthropogenic disturbance on community occupancy.

With biodiversity facing unparalleled threats from anthropogenic disturbance, knowledge on the occurrences of species and communities provides for an effective and fast approach to assess their status and vulnerability. Disturbance is most prominent at the landscape-level, for example through habitat loss from large-scale resource extraction or agriculture. However, addressing species responses to habitat changes at the landscape-scale can be difficult and cost-ineffective, hence studies are mostly conducted at single areas or habitat patches. Moreover, there is a relative lack of studies on communities, as opposed to focal species, despite the former may carry more comprehensive information. Here, we used a multi-region, multi-species hierarchical occupancy model to study a meta-community of mammals detected by camera traps across five distinct areas within a heterogeneous landscape in Tanzania, and aimed to assess responses to human disturbance and environmental variables. Estimated species richness did not vary significantly across different areas, even though these held broadly different habitats. Moreover, we found remarkable consistency in the positive effect of distance to human settlements, a proxy for anthropogenic disturbance, on community occupancy. The positive effect of body size and the positive effect of proximity to rivers on community occupancy were also shared by communities. Results yield conservation relevance because: (1) the among-communities consistency in responses to anthropogenic disturbance, despite the heterogeneity in sampled habitats, indicates that conservation plans designed at the landscape-scale may represent a comprehensive and cost-efficient approach; (2) the consistency in responses to environmental factors suggests that multi-species models are a powerful method to study ecological patterns at the landscape-level.

Reserve size and anthropogenic disturbance affect the density of an African leopard (Panthera pardus) meta-population.

Determining correlates of density for large carnivores is important to understand their ecological requirements and develop conservation strategies. Of several earlier density studies conducted globally, relatively few addressed a scale (usually >1000 km2) that allows inference on correlates of density over heterogeneous landscapes. We deployed 164 camera trap stations covering ~2500 km2 across five areas characterized by broadly different vegetation cover in the Udzungwa Mountains, Tanzania, to investigate correlates of density for a widespread and adaptable carnivore, the leopard (Panthera pardus). We modelled data in a spatially explicit capture-recapture framework, with both biotic and abiotic covariates hypothesised to influence density. We found that leopard density increased with distance to protected area boundary (mean±SE estimated effect = 0.44±0.20), a proxy for both protected area extent and distance from surrounding human settlements. We estimated mean density at 4.22 leopards/100 km2 (85% CI = 3.33‒5.35/100 km2), with no variation across habitat types. Results indicate that protected area extent and anthropogenic disturbance limit leopard populations whereas no support was found for prey availability and trap array as drivers of leopard density. Such vulnerability is relevant to the conservation of the leopard, which is generally considered more resilient to human disturbance than other large cats. Our findings support the notion that protected areas are important to preserve viable population of leopards, increasingly so in times of unprecedented habitat fragmentation. Protection of buffer zones smoothing the abrupt impact of human activities at reserve edges also appears of critical conservation relevance.

A review of the Madagascan pelican spiders of the genera Eriauchenius O. Pickard-Cambridge, 1881 and Madagascarchaea gen. n. (Araneae, Archaeidae).

An endemic genus of Madagascan spiders (Araneae, Archaeidae, Eriauchenius) is revised. All 20 species of Eriauchenius are described and keyed, of which 14 are new species: Eriauchenius andriamanelosp. n., Eriauchenius andrianampoinimerinasp. n., Eriauchenius goodmanisp. n., Eriauchenius harveyisp. n., Eriauchenius lukemacaulayisp. n., Eriauchenius milajaneaesp. n., Eriauchenius millotisp. n., Eriauchenius rafohysp. n., Eriauchenius ranavalonasp. n., Eriauchenius rangitasp. n., Eriauchenius rixisp. n., Eriauchenius samasp. n., Eriauchenius wunderlichisp. n., Eriauchenius zirafysp. n. Additionally, six species of the new genus Madagascarchaeagen. n. are described and keyed, of which four are new species: Madagascarchaea fohysp. n., Madagascarchaea lotzisp. n., Madagascarchaea moramorasp. n., Madagascarchaea rabesahalasp. n. Diagnostic characters for the Madagascan and African genera are described, and based on these characters and previous phylogenetic analyses the following species transfers are proposed: Eriauchenius cornutus (Lotz, 2003) to Afrarchaea; Afrarchaea fisheri (Lotz, 2003) and Afrarchaea mahariraensis (Lotz, 2003) to Eriauchenius. Finally, we propose that the distribution of Afrarchaea be restricted to South Africa. While several Madagascan specimens have previously been identified as Afrarchaea godfreyi (Hewitt, 1919), we argue that these are likely misidentifications that should instead be Eriauchenius.

Gauging megadiversity with optimized and standardized sampling protocols: A case for tropical forest spiders.

Characterizing and monitoring biodiversity and assessing its drivers require accurate and comparable data on species assemblages, which, in turn, should rely on efficient and standardized field collection. Unfortunately, protocols that follow such criteria remain scarce and it is unclear whether they can be applied to megadiverse communities, whose study can be particularly challenging. Here, we develop and evaluate the first optimized and standardized sampling protocol for megadiverse communities, using tropical forest spiders as a model taxon. We designed the protocol COBRA-TF (Conservation Oriented Biodiversity Rapid Assessment for Tropical Forests) using a large dataset of semiquantitative field data from different continents. This protocol combines samples of different collecting methods to obtain as many species as possible with minimum effort (optimized) and widest applicability and comparability (standardized). We ran sampling simulations to assess the efficiency of COBRA-TF (optimized, non-site-specific) and its reliability for estimating taxonomic, phylogenetic, and functional diversity, and community structure by comparing it with (1) commonly used expert-based ad hoc protocols (nonoptimized, site-specific) and (2) optimal protocols (optimized, site-specific). We then tested the performance and feasibility of COBRA-TF in the field. COBRA-TF yielded similar results as ad hoc protocols for species (observed and estimated) and family richness, phylogenetic and functional diversity, and species abundance distribution. Optimal protocols detected more species than COBRA-TF. Data from the field test showed high sampling completeness and yielded low numbers of singletons and doubletons. Optimized and standardized protocols can be as effective in sampling and studying megadiverse communities as traditional sampling, while allowing data comparison. Although our target taxa are spiders, COBRA-TF can be modified to apply to any highly diverse taxon and habitat as long as multiple collecting techniques exist and the unit effort per sample is comparable. Protocols such as COBRA-TF facilitate studying megadiverse communities and therefore may become essential tools for monitoring community changes in space and time, assessing the effects of disturbances and selecting conservation areas.

A new species of Charinus Simon 1892 from Brazil, with notes on behavior (Amblypygi, Charinidae).

A new species of Charinus is described and illustrated from the Brazilian Atlantic Forest. Charinus ruschiisp. n. is found in Santa Lúcia reserve, Espírito Santo state, and is sympatric with Charinus brasilianus and Charinus montanus. The new species can be easily distinguished from the other species of the genus by the combination of the following characters: number of spines on the pedipalp tarsus, size and shape of the female genitalia, shape of the sternum and number of teeth in the cheliceral claw. The behavioral repertory is also described for this species based on five hours of qualitative and 24 hours of quantitative observations in order to define the behavioral categories. Five behavioral categories were detected and 21 behavioral acts. The most conspicuous category was Immobility, followed by Antenniform leg movement, Environmental exploration, Self-grooming, and Feeding. It was also found that juveniles spend longer time inside the shelter, even during peaks of adult activity, which could be related to a survival strategy.

PROBLEMS WITH "SOFT" POLYTOMIES.

Abstract- The "soft" assumption attributes polytomies to lack of data, not simultaneous cladogenesis (the "hard" assumption). Most systematists prefer the first interpretation, but most parsimony programs implicitly use the second. Results can thus be inconsistent with initial assumptions. Under certain circumstances that seem especially typical for large data sets treating higher taxa, it may be valid to eliminate both compatible and incompatible polytomous trees from consideration. Consistent treatment of soft polytomies can reduce the ambiguity of cladistic solutions and improve the resolution, and testability, of phylogenetic hypotheses.

The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling

We present a phylogenetic analysis of spiders using a dataset of 932 spider species, representing 115 families (only the family Synaphridae is unrepresented), 700 known genera, and additional representatives of 26 unidentified or undescribed genera. Eleven genera of the orders Amblypygi, Palpigradi, Schizomida and Uropygi are included as outgroups. The dataset includes six markers from the mitochondrial (12S, 16S, COI) and nuclear (histone H3, 18S, 28S) genomes, and was analysed by multiple methods, including constrained analyses using a highly supported backbone tree from transcriptomic data. We recover most of the higher-level structure of the spider tree with good support, including Mesothelae, Opisthothelae, Mygalomorphae and Araneomorphae. Several of our analyses recover Hypochilidae and Filistatidae as sister groups, as suggested by previous transcriptomic analyses. The Synspermiata are robustly supported, and the families Trogloraptoridae and Caponiidae are found as sister to the Dysderoidea. Our results support the Lost Tracheae clade, including Pholcidae, Tetrablemmidae, Diguetidae, Plectreuridae and the family Pacullidae (restored status) separate from Tetrablemmidae. The Scytodoidea include Ochyroceratidae along with Sicariidae, Scytodidae, Drymusidae and Periegopidae; our results are inconclusive about the separation of these last two families. We did not recover monophyletic Austrochiloidea and Leptonetidae, but our data suggest that both groups are more closely related to the Cylindrical Gland Spigot clade rather than to Synspermiata. Palpimanoidea is not recovered by our analyses, but also not strongly contradicted. We find support for Entelegynae and Oecobioidea (Oecobiidae plus Hersiliidae), and ambiguous placement of cribellate orb-weavers, compatible with their non-monophyly. Nicodamoidea (Nicodamidae plus Megadictynidae) and Araneoidea composition and relationships are consistent with recent analyses. We did not obtain resolution for the titanoecoids (Titanoecidae and Phyxelididae), but the Retrolateral Tibial Apophysis clade is well supported. Penestomidae, and probably Homalonychidae, are part of Zodarioidea, although the latter family was set apart by recent transcriptomic analyses. Our data support a large group that we call the marronoid clade (including the families Amaurobiidae, Desidae, Dictynidae, Hahniidae, Stiphidiidae, Agelenidae and Toxopidae). The circumscription of most marronoid families is redefined here. Amaurobiidae include the Amaurobiinae and provisionally Macrobuninae. We transfer Malenellinae (Malenella, from Anyphaenidae), Chummidae (Chumma) (new syn.) and Tasmarubriinae (Tasmarubrius, Tasmabrochus and Teeatta, from Amphinectidae) to Macrobuninae. Cybaeidae are redefined to include Calymmaria, Cryphoeca, Ethobuella and Willisius (transferred from Hahniidae), and Blabomma and Yorima (transferred from Dictynidae). Cycloctenidae are redefined to include Orepukia (transferred from Agelenidae) and Pakeha and Paravoca (transferred from Amaurobiidae). Desidae are redefined to include five subfamilies: Amphinectinae, with Amphinecta, Mamoea, Maniho, Paramamoea and Rangitata (transferred from Amphinectidae); Ischaleinae, with Bakala and Manjala (transferred from Amaurobiidae) and Ischalea (transferred from Stiphidiidae); Metaltellinae, with Austmusia, Buyina, Calacadia, Cunnawarra, Jalkaraburra, Keera, Magua, Metaltella, Penaoola and Quemusia; Porteriinae (new rank), with Baiami, Cambridgea, Corasoides and Nanocambridgea (transferred from Stiphidiidae); and Desinae, with Desis, and provisionally Poaka (transferred from Amaurobiidae) and Barahna (transferred from Stiphidiidae). Argyroneta is transferred from Cybaeidae to Dictynidae. Cicurina is transferred from Dictynidae to Hahniidae. The genera Neoramia (from Agelenidae) and Aorangia, Marplesia and Neolana (from Amphinectidae) are transferred to Stiphidiidae. The family Toxopidae (restored status) includes two subfamilies: Myroinae, with Gasparia, Gohia, Hulua, Neomyro, Myro, Ommatauxesis and Otagoa (transferred from Desidae); and Toxopinae, with Midgee and Jamara, formerly Midgeeinae, new syn. (transferred from Amaurobiidae) and Hapona, Laestrygones, Lamina, Toxops and Toxopsoides (transferred from Desidae). We obtain a monophyletic Oval Calamistrum clade and Dionycha; Sparassidae, however, are not dionychans, but probably the sister group of those two clades. The composition of the Oval Calamistrum clade is confirmed (including Zoropsidae, Udubidae, Ctenidae, Oxyopidae, Senoculidae, Pisauridae, Trechaleidae, Lycosidae, Psechridae and Thomisidae), affirming previous findings on the uncertain relationships of the "ctenids" Ancylometes and Cupiennius, although a core group of Ctenidae are well supported. Our data were ambiguous as to the monophyly of Oxyopidae. In Dionycha, we found a first split of core Prodidomidae, excluding the Australian Molycriinae, which fall distantly from core prodidomids, among gnaphosoids. The rest of the dionychans form two main groups, Dionycha part A and part B. The former includes much of the Oblique Median Tapetum clade (Trochanteriidae, Gnaphosidae, Gallieniellidae, Phrurolithidae, Trachelidae, Gnaphosidae, Ammoxenidae, Lamponidae and the Molycriinae), and also Anyphaenidae and Clubionidae. Orthobula is transferred from Phrurolithidae to Trachelidae. Our data did not allow for complete resolution for the gnaphosoid families. Dionycha part B includes the families Salticidae, Eutichuridae, Miturgidae, Philodromidae, Viridasiidae, Selenopidae, Corinnidae and Xenoctenidae (new fam., including Xenoctenus, Paravulsor and Odo, transferred from Miturgidae, as well as Incasoctenus from Ctenidae). We confirm the inclusion of Zora (formerly Zoridae) within Miturgidae.

Why do tropical mountains support exceptionally high biodiversity? The Eastern Arc mountains and the drivers of Saintpaulia diversity.

We combine information about the evolutionary history and distributional patterns of the genus Saintpaulia H. Wendl. (Gesneriaceae; 'African violets') to elucidate the factors and processes behind the accumulation of species in tropical montane areas of high biodiversity concentration. We find that high levels of biodiversity in the Eastern Arc Mountains are the result of pre-Quaternary speciation processes and environmental stability. Our results support the hypothesis that climatically stable mountaintops may have acted as climatic refugia for lowland lineages during the Pleistocene by preventing extinctions. In addition, we found evidence for the existence of lowland micro-refugia during the Pleistocene, which may explain the high species diversity of East African coastal forests. We discuss the conservation implications of the results in the context of future climate change.

The velvet spiders: an atlas of the Eresidae (Arachnida, Araneae).

The family Eresidae C. L. Koch, 1850 is reviewed at the genus level. The family comprises nine genera including one new genus. They are: Adonea Simon, 1873, Dorceus C. L. Koch, 1846, Dresserus Simon, 1876, Eresus Walckenaer, 1805, Gandanameno Lehtinen, 1967, Loureediagen. n., ParadoneaLawrence, 1968, Seothyra Purcell, 1903, and Stegodyphus Simon, 1873. A key to all genera and major lineages is provided along with corresponding diagnoses, as well as descriptions of selected species. These are documented with collections of photographs, scanning electron micrographs, and illustrations. A new phylogeny of Eresidae based on molecular sequence data expands on a previously published analysis. A species of the genus Paradonea Lawrence, 1968 is sequenced and placed phylogenetically for the first time. New sequences from twenty Gandanameno Lehtinen, 1967 specimens were added to investigate species limits within the genus. The genus Loureediagen. n. is proposed to accommodate Eresus annulipes Lucas, 1857. Two species, Eresus semicanus Simon, 1908 and Eresus jerbae El-Hennawy, 2005, are synonymized with Loureedia annulipescomb. n. One new species, Paradonea presleyisp. n. is described. Eresus algericus El-Hennawy, 2004 is transferred to Adonea Simon, 1873. The female of Dorceus fastuosus C. L. Koch, 1846 is described for the first time. The first figures depicting Paradonea splendens (Lawrence, 1936) are presented.