Daniele Polotow

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.

Total evidence analysis of the phylogenetic relationships of Lycosoidea spiders (Araneae, Entelegynae)

Abstract. Phylogenetic relationships within the superfamily Lycosoidea are investigated through the coding and analysis of character data derived from morphology, behaviour and DNA sequences. In total, 61 terminal taxa were studied, representing most of the major groups of the RTA-clade (i.e. spiders that have a retrolateral tibial apophysis on the male palp). Parsimony and model-based approaches were used, and several support values, partitions and implied weighting schemes were explored to assess clade stability. The morphological–behavioural matrix comprised 96 characters, and four gene fragments were used: 28S (∼737 base pairs), actin (∼371 base pairs), COI (∼630 base pairs) and H3 (∼354 base pairs). Major conclusions of the phylogenetic analysis include: the concept of Lycosoidea is restricted to seven families: Lycosidae, Pisauridae, Ctenidae, Psechridae, Thomisidae, Oxyopidae (but Ctenidae and Pisauridae are not monophyletic) and also Trechaleidae (not included in the analysis); the monophyly of the ‘Oval Calamistrum clade’ (OC-clade) appears to be unequivocal, with high support, and encompassing the Lycosoidea plus the relimited Zoropsidae and the proposed new family Udubidae (fam. nov.); Zoropsidae is considered as senior synonym of Tengellidae and Zorocratidae (syn. nov.); Viridasiinae (rank nov.) is raised from subfamily to family rank, excluded from the Ctenidae and placed in Dionycha. Our quantitative phylogenetic analysis confirms the synonymy of Halidae with Pisauridae. The grate-shaped tapetum appears independently at least three times and has a complex evolutionary history, with several reversions.

Description of Toca, a new neotropical spider genus (Araneae, Ctenidae, Calocteninae)

Abstract Toca new genus is proposed to include two new species: the type species T. bossanova new species from Rio de Janeiro, Brazil, and T. samba new species from Paraná and Minas Gerais, Brazil. Toca may be related to Caloctenus Keyserling and Gephyroctenus Mello-Leitão, with which it shares the scales on the abdominal dorsum and the epigynum as a single, slightly sclerotized, fold. The genus can be distinguished among the Calocteninae genera by its unique genital structures.

Phylogenetic relationships of the Neotropical spider genus Itatiaya (Araneae)

Polotow, D. & Brescovit, A. D. (2010). Phylogenetic relationships of the Neotropical spider genus Itatiaya (Araneae). —Zoologica Scripta, 40, 187–193.

The new spider genus Arctenus, an afrotropical representative of the Calocteninae (Araneae: Ctenidae)

Abstract Arctenus gen. nov. is proposed to include the type species A. taitensis sp. nov. from the Taita Hills in Kenya. This ctenid species appears to be the first representative of the Calocteninae in the African continent. Results of a parsimony analysis of morphological and behavioral characters indicated that the new species cannot be placed in any known genus and therefore validated the creation of the new genus whose autapomorphies are considered hypotheses for the genus synapomorphies. The phylogenetic relationships of the new genus are discussed and a distribution map of the unique species is presented.

Redescription of five Amazonian species of Ctenus Walckenaer (Araneae, Ctenidae, Cteninae)

Five species of Ctenus from the Amazon basin are redescribed: C. delesserti (Caporiacco, 1947), C. falconensis Schenkel, 1953, C. nigritus F.O. Pickard-Cambridge, 1897, C. serratipes F.O. Pickard-Cambridge, 1897 and C. sigma (Schenkel, 1953). Three new synonymies are proposed: Ctenus fulvipes Caporiacco, 1947, C. itatiayaeformis Caporiacco, 1955 and C. scenicus Caporiacco, 1947 with C. serratipes. The female of Ctenus nigritus is described for the first time. The distributional ranges of Ctenus nigritus and C. serratipes are extended. We also present distributional maps of the five redescribed species.

Systematics and biogeography of Spinoctenus, a new genus of wandering spider from Colombia (Ctenidae)

Abstract. Among ctenid spiders, ctenines comprise the most diverse subfamily. In this study, a new genus of Cteninae, Spinoctenus, is proposed to include the type species S. yotoco, sp. nov. Ten new species are also described: S. escalerete, S. pericos, S. eberhardi, S. spinosus, S. stephaniae, S. nambi, S. florezi, S. tequendama, S. chocoensis and S. flammigerus. Results of the parsimony and Bayesian phylogenetic analyses using morphological and behavioural characters indicate the monophyly of this genus, closely related to Phoneutria Perty, 1883 and Ctenus Walckenaer, 1805. This genus can be distinguished from the remaining Ctenidae by three unambiguous synapomorphies: embolus with folded process, tegulum with median process, and RTA curved internally close to the cymbium. A dispersal-vicariance biogeographical analysis of the genus in the Andean and Chocó regions indicates the origin of Spinoctenus in the Western and Central Andean Cordilleras. From this region, three events of dispersal occurred to the other regions (one to the Chocó and two to the Eastern Cordillera), which were subsequently followed by three events of vicariance, suggesting that dispersal and vicariance were equally important in shaping the current distribution patterns of Spinoctenus species. The discovery of this new genus containing a large number of new species in the Andean and Chocó regions highlights the current poor knowledge of the Colombian biodiversity.