The infraorder Mygalomorphae is one of the three main lineages of spiders comprising over 3000 nominal species. This ancient group has a worldwide distribution that includes among its ranks large and charismatic taxa such as tarantulas, trapdoor spiders, and highly venomous funnel-web spiders. Based on past molecular studies using Sanger-sequencing approaches, numerous mygalomorph families (e.g., Hexathelidae, Ctenizidae, Cyrtaucheniidae, Dipluridae, and Nemesiidae) have been identified as non-monophyletic. However, these data were unable to sufficiently resolve the higher-level (intra- and interfamilial) relationships such that the necessary changes in classification could be made with confidence. Here, we present a comprehensive phylogenomic treatment of the spider infraorder Mygalomorphae. We employ 472 loci obtained through anchored hybrid enrichment to reconstruct relationships among all the mygalomorph spider families and estimate the timeframe of their diversification. We sampled nearly all currently recognized families, which has allowed us to assess their status, and as a result, propose a new classification scheme. Our generic-level sampling has also provided an evolutionary framework for revisiting questions regarding silk use in mygalomorph spiders. The first such analysis for the group within a strict phylogenetic framework shows that a sheet web is likely the plesiomorphic condition for mygalomorphs, as well as providing insights to the ancestral foraging behavior for all spiders. Our divergence time estimates, concomitant with detailed biogeographic analysis, suggest that both ancient continental-level vicariance and more recent dispersal events have played an important role in shaping modern day distributional patterns. Based on our results, we relimit the generic composition of the Ctenizidae, Cyrtaucheniidae, Dipluridae, and Nemesiidae. We also elevate five subfamilies to family rank: Anamidae (NEW RANK), Euagridae (NEW RANK), Ischnothelidae (NEW RANK), Pycnothelidae (NEW RANK), and Bemmeridae (NEW RANK). Three families Entypesidae (NEW FAMILY), Microhexuridae (NEW FAMILY), and Stasimopidae (NEW FAMILY), and one subfamily Australothelinae (NEW SUBFAMILY) are newly proposed. Such a major rearrangement in classification, recognizing nine newly established family-level rank taxa, is the largest the group has seen in over three decades. [Biogeography; molecular clocks; phylogenomics; spider web foraging; taxonomy.]
Abstract. The aim of this study was to characterize karyotypes of central European spiders of the genera Arctosa, Tricca, and Xerolycosa (Lycosidae) with respect to the diploid chromosome number, chromosome morphology, and sex chromosomes. Karyotype data are reported for eleven species, six of them for the first time. For selected species the pattern in the distributions of the constitutive heterochromatin and the nucleolar organizer regions (NORs) was determined. The silver staining technique for detecting NORs of lycosid spiders was standardized. The male karyotype consisted of 2n = 28 (Arctosa and Tricca) or 2n = 22 (Xerolycosa) acrocentric chromosomes. The sex chromosome system was X1X20 in all species. The sex chromosomes of T. lutetiana and X. nemoralis showed unusual behaviour during late diplotene, namely temporary extension due to decondensation. C-banding technique revealed a small amount of constitutive heterochromatin at the centromeric region of the chromosomes. Two pairs of autosomes bore terminal NORs. Differences in karyotypes among Arctosa species indicate that the evolution of the karyotype in this genus involved autosome translocations and size changes in the sex chromosomes. Based on published results and those recorded in this study it is suggested that the ancestral male karyotype of the superfamily Lycosoidea consisted of 28 acrocentric chromosomes.
Studies conducted on volcanic islands have greatly contributed to our current understanding of how organisms diversify. The Canary Islands archipelago, located northwest of the coast of northern Africa, harbours a large number of endemic taxa. Because of their low vagility, mygalomorph spiders are usually absent from oceanic islands. The spider Titanidiops canariensis, which inhabits the easternmost islands of the archipelago, constitutes an exception to this rule. Here, we use a multi-locus approach that combines three mitochondrial and four nuclear genes to investigate the origins and phylogeography of this remarkable trap-door spider. We provide a timeframe for the colonisation of the Canary Islands using two alternative approaches: concatenation and species tree inference in a Bayesian relaxed clock framework. Additionally, we investigate the existence of cryptic species on the islands by means of a Bayesian multi-locus species delimitation method. Our results indicate that T. canariensis colonised the Canary Islands once, most likely during the Miocene, although discrepancies between the timeframes from different approaches make the exact timing uncertain. A complex evolutionary history for the species in the archipelago is revealed, which involves two independent colonisations of Fuerteventura from the ancestral range of T. canariensis in northern Lanzarote and a possible back colonisation of southern Lanzarote. The data further corroborate a previously proposed volcanic refugium, highlighting the impact of the dynamic volcanic history of the island on the phylogeographic patterns of the endemic taxa. T. canariensis includes at least two different species, one inhabiting the Jandia peninsula and central Fuerteventura and one spanning from central Fuerteventura to Lanzarote. Our data suggest that the extant northern African Titanidiops lineages may have expanded to the region after the islands were colonised and, hence, are not the source of colonisation. In addition, T. maroccanus may harbour several cryptic species.
BackgroundA large scale semi-quantitative biodiversity assessment was conducted in white oak woodlands in areas included in the Spanish Network of National Parks, as part of a project aimed at revealing biogeographic patterns and identify biodiversity drivers. The semi-quantitative COBRA sampling protocol was conducted in sixteen 1-ha plots across six national parks using a nested design. All adult specimens were identified to species level based on morphology. Uncertain delimitations and identifications due to either limited information of diagnostic characters or conflicting taxonomy were further investigated using DNA barcode information.New informationWe identified 376 species belonging to 190 genera in 39 families, from the 8,521 adults found amongst the 20,539 collected specimens. Faunistic results include the discovery of 7 new species to the Iberian Peninsula, 3 new species to Spain and 11 putative new species to science. As largely expected by environmental features, the southern parks showed a higher proportion of Iberian and Mediterranean species than the northern parks, where the Palearctic elements were largely dominant. The analysis of approximately 3,200 DNA barcodes generated in the present study, corroborated and provided finer resolution to the morphologically based delimitation and identification of specimens in some taxonomically challenging families. Specifically, molecular data confirmed putative new species with diagnosable morphology, identified overlooked lineages that may constitute new species, confirmed assignment of specimens of unknown sexes to species and identified cases of misidentifications and phenotypic polymorphisms.
The family Hexathelidae ranks among the smaller mygalomorph spider families. Most species are endemic to the Australasian region and the family was traditionally considered an example of a Gondwanan lineage. However, recent studies have cast some doubt on the monophyly of the family. Macrothele is the only genus with an out-of-Gondwana distribution. The bulk of the Macrothele diversity is found in South-east Asia, few species are known from central Africa and two species inhabit Europe: Macrothele calpeiana (Walckenaer, 1805) from the Iberian Peninsula and Macrothele cretica Kulczynski, 1903 endemic to Crete. Here we investigate the origins of the European Macrothele species by means of a multi-locus phylogenetic approach and by inferring the time frame of the diversification of the genus using Bayesian relaxed clock methods. We also provide further insights into the phylogenetic status of the family Hexathelidae. Our results indicate that the diversification of Macrothele traces back to the period of the Gondwana break-up and its present-day distribution most likely reflects the subsequent tectonic plate movements. The two European species were not recovered as sister taxa, suggesting that Macrothele colonised the Mediterranean region twice independently. The polyphyly of the family Hexathelidae is further confirmed and the subfamily Atracinae is identified as the conflicting lineage.
26The Infraorder Mygalomorphae is one of the three main lineages of spiders comprising over 27 3,000 nominal species. This ancient group has a world-wide distribution that includes among its 28 ranks large and charismatic taxa such as tarantulas, trapdoor spiders, and highly venomous 29 funnel web spiders. Based on past molecular studies using Sanger-sequencing approaches, 30 numerous mygalomorph families (e.g., Hexathelidae, Ctenizidae, Cyrtaucheniidae, Dipluridae 31 and Nemesiidae) have been identified as non-monophyletic. However, these data were unable to 32 sufficiently resolve the higher-level (intra-and interfamilial) relationships such that the 33 necessary changes in classification could be made with confidence. Here we present a 34 comprehensive phylogenomic treatment of the spider infraorder Mygalomorphae. We employ 35 472 loci obtained through Anchored Hybrid Enrichment to reconstruct relationships among all 36 the mygalomorph spider families and estimate the timeframe of their diversification. We 37 sampled all currently recognized families, which has allowed us to assess their status, and as a 38 result, propose a new classification scheme. Our generic-level sampling has also provided an 39 evolutionary framework for revisiting questions regarding silk use in mygalomorph spiders. The 40 first such analysis for the group within a strict phylogenetic framework shows that a sheet web is 41 likely the plesiomorphic condition for mygalomorphs, as well as providing hints to the ancestral 42 foraging behavior for all spiders. Our divergence time estimates, concomitant with detailed 43 biogeographic analysis, suggest that both ancient continental-level vicariance and more recent 44 dispersal events have played an important role in shaping modern day distributional patterns. 45Based on our results, we relimit the generic composition of the Ctenizidae, Cyrtaucheniidae, 46Dipluridae and Nemesiidae. We also elevate five subfamilies to family rank: Anamidae (NEW 47 PHYLOGENY OF MYGALOMORPH SPIDERS 3 RANK), Euagridae (NEW RANK), Ischnothelidae (NEW RANK), Pycnothelidae (NEW 48 RANK), and Bemmeridae (NEW RANK). The three families Entypesidae (NEW FAMILY), 49 Microhexuridae (NEW FAMILY), and Stasimopidae (NEW FAMILY) are newly proposed. 50 Such a major rearrangement in classification, recognizing eight newly established family-level 51 rank taxa, is the largest the group has seen in over three decades since Raven's (1985) taxonomic 52 treatment. 53 54
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