The spider tree of life: phylogeny of Araneae based on target‐gene analyses from an extensive taxon sampling
Authors: Wheeler Ward C. , Coddington Jonathan A. , Crowley Louise M. , Dimitrov Dimitar , Goloboff Pablo A. , Griswold Charles E. , Hormiga Gustavo , Prendini Lorenzo , Ramírez Martín J. , Sierwald Petra , Almeida‐Silva Lina , Alvarez‐Padilla Fernando , Arnedo Miquel A. , Benavides Silva Ligia R. , Benjamin Suresh P. , Bond Jason E. , Grismado Cristian J. , Hasan Emile , Hedin Marshal , Izquierdo Matías A. , Labarque Facundo M. , Ledford Joel , Lopardo Lara , Maddison Wayne P. , Miller Jeremy A. , Piacentini Luis N. , Platnick Norman I. , Polotow Daniele , Silva‐Dávila Diana , Scharff Nikolaj , Szűts Tamás , Ubick Darrell , Vink Cor J. , Wood Hannah M. , Zhang Junxia 
Published: December 2016

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,TasmabrochusandTeeatta, from Amphinectidae) to Macrobuninae. Cybaeidae are redefined to includeCalymmaria,Cryphoeca,EthobuellaandWillisius(transferred from Hahniidae), andBlabommaandYorima(transferred from Dictynidae). Cycloctenidae are redefined to includeOrepukia(transferred from Agelenidae) andPakehaandParavoca(transferred from Amaurobiidae). Desidae are redefined to include five subfamilies: Amphinectinae, withAmphinecta,Mamoea,Maniho,ParamamoeaandRangitata(transferred from Amphinectidae); Ischaleinae, withBakalaandManjala(transferred from Amaurobiidae) andIschalea(transferred from Stiphidiidae); Metaltellinae, withAustmusia,Buyina,Calacadia,Cunnawarra,Jalkaraburra,Keera,Magua,Metaltella,PenaoolaandQuemusia; Porteriinae (new rank), withBaiami,Cambridgea,CorasoidesandNanocambridgea(transferred from Stiphidiidae); and Desinae, withDesis, and provisionallyPoaka(transferred from Amaurobiidae) andBarahna(transferred from Stiphidiidae).Argyronetais transferred from Cybaeidae to Dictynidae.Cicurinais transferred from Dictynidae to Hahniidae. The generaNeoramia(from Agelenidae) andAorangia,MarplesiaandNeolana(from Amphinectidae) are transferred to Stiphidiidae. The family Toxopidae (restored status) includes two subfamilies: Myroinae, withGasparia,Gohia,Hulua,Neomyro,Myro,OmmatauxesisandOtagoa(transferred from Desidae); and Toxopinae, withMidgeeandJamara, formerly Midgeeinae,new syn.(transferred from Amaurobiidae) andHapona,Laestrygones,Lamina,ToxopsandToxopsoides(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”AncylometesandCupiennius, 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.Orthobulais 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., includingXenoctenus,ParavulsorandOdo, transferred from Miturgidae, as well asIncasoctenusfrom Ctenidae). We confirm the inclusion ofZora(formerly Zoridae) within Miturgidae.