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 rede...
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.
The monophyly of Tetragnathidae including the species composition of the family (e.g., Are Nephila and their relatives part of this lineage?), the phylogenetic relationships of its various lineages, and the exact placement of Tetragnathidae within Araneoidea have been three recalcitrant problems in spider systematics. Most studies on tetragnathid phylogeny have focused on morphological and behavioral data, but little molecular work has been published to date. To address these issues we combine previous morphological and behavioral data with novel molecular data including nuclear ribosomal RNA genes 18S and 28S, mitochondrial ribosomal RNA genes 12S and 16S and protein-coding genes from the mitochondrion [cytochrome c oxidase subunit I (COI)] and from the nucleus (histone H3), totaling ca. 6.3 kb of sequence data per taxon. These data were analyzed using direct optimization and static homology using both parsimony and Bayesian methods. Our results indicate monophyly of Tetragnathidae, Tetragnathinae, Leucauginae, the ''Nanometa clade'' and the subfamily Metainae, which, with the exception of the later subfamily, received high nodal support. Morphological synapomorphies that support these clades are also discussed. The position of tetragnathids with respect to the rest of the araneoid spiders remains largely unresolved but tetragnathids and nephilids were never recovered as sister taxa. The combined dataset suggests that Nephilidae is sister to Araneidae; furthermore, the sister group of Nephila is the clade composed by Herennia plus Nephilengys and this pattern has clear implications for understanding the comparative biology of the group. Tetragnathidae is most likely sister to some members of the ''reduced piriform clade'' and nephilids constitute the most-basal lineage of araneids.
The spider genus Metabus (Tetragnathidae) previously included nine species: the type M. gravidus O. P.-Cambridge, 1899 -junior synonym of Leucauge ocellata (Keyserling) -from Central America and eight species from Chile. In this paper, the classification of the Metabus species-complex is revised, and two new genera, with three new species and five new combinations, are described. Allende gen. nov. is created for four Chilean species not congeneric with the type of Metabus : the type A. puyehuensis sp. nov. , A. patagiatus (Simon) comb. nov. , A. nigrohumeralis (F. O. P.-Cambridge) comb. nov. and A. longipes (Nicolet) comb. nov. Further additions to the Chilean fauna are under the new genus Mollemeta gen. nov. -created for M. edwardsi (Simon) comb. nov. -and three new species of Chrysometa : C. acinosa sp. nov. , C. levii sp. nov. and C. maitae sp. nov. Metabus now includes four species: M. ocellatus (Keyserling) comb. nov. , M. debilis (O. P.-Cambridge) comb. nov. , M. ebanoverde sp. nov. and M.conacyt sp. nov. All of these species were included in a phylogenetic analysis of 38 tetragnathid and 12 orbicularian outgroup terminals scored for 105 morphological and behavioural characters. The results suggest that Metabus as previously circumscribed is polyphyletic. The phylogenetic relationships within tetragnathids are briefly discussed.
The present atlas documents the morphology of representative species of 22 tetragnathid genera, with emphasis on nontetragnathines. It includes approximately 960 scanning electron micrographs, morphological drawings and web photographs. The 213 characters used in the phylogenetic analyses are described and illustrated. We discuss the optimal cladograms obtained by the analysis of the morphological and behavioural data, and compare our results to a recent hypothesis of tetragnathid phylogenetic relationships that combined similar data with multigene DNA sequences. Based on the cladistic hypothesis that results from the total evidence analysis, we study the evolution of six morphological character systems within Tetragnathidae: spinneret spigots, respiratory structures, trichobothria, chelicerae, and male and female genitalia.
The female genital morphology of the spiders in the araneoid genus Agriognatha (Tetragnathidae) is described and illustrated. The female genitalia of Agriognatha is characterized by a strong reduction of the sperm storage organs (spermathecae) and by the presence of a specialized distal compartment of the median membranous chamber that functions as a sperm storage organ (the posterior sac). The genital morphology of Agriognatha species is unique among Tetragnathidae and it provides robust synapomorphic evidence for the monophyly of genus. We discuss the phylogenetic implications of these new findings for the placement and monophyly of Agriognatha and for the monophyly of Tetragnathinae.
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