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.

New species of the neotropical spider genus Celaetycheus Simon, 1897 (Araneae: Ctenidae).

Nine species of the Neotropical spider genus Celaetycheus Simon, 1897 are newly described. All the species are endemic to the State of Bahia, northeast of Brazil: Celaetycheus abara sp. nov., C. aberen sp. nov., C. acaraje sp. nov., C. beiju sp. nov., C. bobo sp. nov., C. caruru sp. nov., C. moqueca sp. nov., C. mungunza sp. nov. and C. vatapa sp. nov. A new generic diagnosis, species diagnoses and distributional maps are provided.

New records and geographical distribution of ctenid spiders (Araneae: Ctenidae) in Colombia.

This study provides new records, geographical distribution extensions and a checklist of the current ctenids species in Colombia based on the review of four arachnological collections and published literature. A total of 15 new records for Ctenidae in Colombia are reported; nine of these species are new records for the country and the distribution of the remaining six is expanded. The genus Centroctenus Mello-Leitão, 1929 (C. ocelliventer Strand, 1909) is recorded for first time in Colombia and Cupiennius coccineus (F.O. Pickard-Cambridge, 1901) for South America. Due to the strategic geographic position of Colombia, which is a transition zone between Southern and Central American biotas, species inventories in different localities are important to fill distributional gaps. The number of known species of ctenids in Colombia is increased from 16 to 25 and these data will be useful for future studies in taxonomy, systematics and biogeography of this family.

Description of three new species of Isoctenus Bertkau, 1880 and new records for I. areia Polotow & Brescovit (Araneae, Ctenidae) from northeastern Brazil.

Three new species of the genus Isoctenus Bertkau, 1880 are described from the state of Bahia, in the northeast of Brazil: Isoctenus jussara new species, Isoctenus cabula new species, and Isoctenus botocudos new species. In addition, new records of Isoctenus areia Polotow and Brescovit, 2009 are presented. Previously known only from the city of Areia, state of Paraíba, its distribution now expands to the cities of Mamanguape, state of Paraíba, and Igarassu, state of Pernambuco.

Revision of the Neotropical spider genus Acanthoctenus (Araneae: Ctenidae: Acanthocteninae).

We revise the genus Acanthoctenus Keyserling, 1877 recognizing thirteen valid species, of which five are new species and two are re-validated. Further, we find one new synonymy and transfer one species. We describe Acanthoctenus alux sp. nov. from Guatemala, A. chickeringi sp. nov. and A. lamarrei sp. nov. from Panama, A. manauara sp. nov. from Brazil and A. torotoro sp. nov. from Bolivia. We revalidate Acanthoctenus dumicola Simon, 1906 stat. res. from Venezuela, and A. virginea (Kraus, 1955) stat. res., comb. nov. from El Salvador. We transfer Acanthoctenus mammifer to Viracucha mammifer (Mello-Leitão, 1939) comb. nov., from Brazil. Acanthoctenus maculatus Petrunkevitch, 1925 and Gephyroctenus kolosvaryi Caporiacco, 1947 are considered species inquirendae in Acanthocteninae and Ctenidae, respectively, and A. obauratus Simon, 1906 and A. rubrotaeniatus Mello-Leitão, 1947 are considered incertae sedis in Acanthocteninae and Acantheinae, respectively. We also describe for the first time the female of Acanthoctenus spiniger Keyserling, 1877, the type species of the genus. We provide illustrations of male and female diagnostic characters, genitalia, habitus, and measurements to support the genus re-description and further identification of its species. We yield a distributional map of the specimens recorded and the description of the natural history of Acanthoctenus manauara sp. nov.

Description of a new Neotropical spider genus in the family Ctenidae Keyserling, 1877 (Araneae: Lycosoidea) from the Brazilian Amazon rainforest.

A new Neotropical genus of Ctenidae from the Brazilian Amazon rainforest is described based on male and female characters: Bulboctenus gen. nov. Three new species are described for the genus: Bulboctenus kayapo sp. nov., Bulboctenus itunaitata sp. nov. and Bulboctenus munduruku sp. nov. The genus can be distinguished from other Ctenidae by males with median apophysis oriented horizontally, perpendicular to the tegulum, RTA strongly excavated anteriorly, metatarsi III and IV with bulbous setae ventrally, sternum and coxae with thick setae; and females with median sector of the epigyne hexagonal with two posterior conical projections ventrally and elevated lateral margins. Discussion on the modification on the leg setae across the Neotropical Ctenids species are given.

Six new species of the spider genus Centroctenus Mello-Leitão from the Neotropical region (Ctenidae, Cteninae).

The species of the genus Centroctenus can be distinguished from other Ctenidae genera by the presence of a long tibiae and the absence of a retrolateral projection of the cymbium in the male palp, and by the presence of a rounded spermathecae in the female epigynum. The composition of this spider genus is herein expanded to eleven Neotropical species, with the description of six new species: Centroctenus dourados sp. n., C. claudia sp. n., C. chalkidisi sp. n., and C. varzea sp. n., from Brazil; and Centroctenus coloso sp. n. from Colombia and Centroctenus alinahui sp. n. from Ecuador. A map with the distribution of all known species in the genus is presented.

Almost eight decades later: description of the male of Isoctenus guadalupei (Mello-Leitão, 1941) comb. nov. and redescription of the female.

Isoctenus Bertkau 1880 is a genus composed by medium to large sized (7 to 19 mm of total body length) Neotropical spiders (Polotow Brescovit 2009). They are nocturnal wandering hunters, which prey in the ground and above leaf litter, as the other tropical wolf spiders of Ctenidae (Griswold et al. 2005). The genus was revised by Polotow Brescovit (2009) and currently contains 15 species, distributed in the Parana (Atlantic, Parana, and Araucaria Forests provinces) and Chacoan (Cerrado and Pampean provinces) dominions (Morrone 2014). Polotow Brescovit (2009) also presented a phylogenetic analysis based on morphological characters, supporting the transference of four species from Ctenus Walckenaer, 1805 to Isoctenus. Later, the genus was included by Polotow Brescovit (2014) in a phylogenetic analysis of Cteninae, and it appears to be related to the Neotropical Parabatinga Polotow Brescovit, 2009 and the Afrotropical Petaloctenus Jocqué Steyn, 1997 and Africactenus Hyatt, 1954, in addition to a couple of misplaced Ctenus species.

Guasuctenus gen. nov., a new Neotropical spider genus of Ctenidae (Araneae).

The new genus Guasuctenus gen. nov. is described based on the results of a recent phylogenetic analysis. This phylogeny, which included representatives of all Ctenidae subfamilies, showed that the genus Ctenus as currently defined, does not comprise a monophyletic group. The results indicated that several species are misplaced in Ctenus and should be transferred to new genera. Aiming to correct the placement of two species originally described in Ctenus, we propose the new genus Guasuctenus to accommodate those species and better represent their phylogenetic relationships. Guasuctenus is supported by the presence of a dorsal cymbial projection in the male palp. Previously regarded as a subspecies, Ctenus longipes vittatissimus Strand, 1916 is here ranked as species, and Guasuctenus longipes (Keyserling, 1891) new comb. and G. vittatissimus (Strand, 1916) new rank, new comb. are redescribed. Ctenus griseus Keyserling, 1891 is recognized as a junior synyonym of Ctenus longipes Keyserling, 1891. Lectotypes and paralectotypes of Guasuctenus vittatissimus (Strand, 1016) are designated for stability. The species of the genus are distributed in southern Brazil and Uruguay.

Chinja, a new genus of spider from the Eastern Arc Mountains of Tanzania (Araneae, Zoropsidae).

The new zoropsid spider genus Chinja Polotow Griswold is diagnosed and described and the following two new species are described: C. chinja sp. nov. and C. scharffi sp. nov. The species were collected in the Eastern Arc Mountains of Tanzania and appear to be endemic to this region. The genus belongs to the oval-calamistrum clade (OC Clade), which also includes Dionycha and Lycosoidea. Based on the synapomorphies and diagnostic characters of members of the OC Clade, Chinja is considered a member of Zoropsidae, although without a clear subfamily placement due to insufficient data. Males of Chinja can be distinguished from other Zoropsidae by a straight PER, by lacking a tibial crack, by having a male palpal cymbium with a retrobasal process and the male palpal tibia with an RTA and a retromedian cluster of stout setae. Females have a divided cribellum with cribellar spigots evenly arranged, and the epigynum with the median plate broad and laterally procurved into hooks, and the lateral lobes each with a wide tooth. The following set of characters can be also helpful to identify the genus: presence of a third tarsal claw, absence of claw tufts and presence of a cribellum and calamistrum.

Kiekie, a new Neotropical spider genus of Ctenidae (Cteninae, Araneae).

Kiekie gen. nov. is newly proposed to accommodate 11 Neotropical Ctenidae species. With the exception of one species from Colombia, they are all distributed in Central America. Two species are transferred from Ctenus: Kiekie sinuatipes (F.O. Pickard-Cambridge, 1897) comb. nov. from Costa Rica, and K. curvipes (Keyserling, 1881) comb. nov., with broader distribution from Mexico to Panama. Ctenus incolans (F.O. Pickard-Cambridge, 1900) is synonymized with Kiekie curvipes. Nine species are newly described. Kiekie garifuna sp. nov. was collected in Guatemala and Honduras. Four species were collected in Costa Rica: Kiekie sanjose sp. nov., K. verbena sp. nov., K. sarapiqui sp. nov., and K. griswoldi sp. nov.. Two species were collected in Panama: Kiekie barrocolorado sp. nov. and K. panamensis sp. nov.. Kiekie montanensis sp. nov. was collected in Costa Rica and Panama. Kiekie antioquia sp. nov., from Colombia, was the only species collected in South America. To support the description and further identification of the genus and its species, we provide illustrations of male and female genitalia, habitus and modified metatarsus of the male leg IV, measurements and diagnostic characters. We also provide a distributional map of the specimens herein recorded. Taxonomic changes are based in recent cladistics analysis, showing that at least two species belong to a well-supported clade, sister group of Africactenus Hyatt, 1954. We discuss the phylogenetic relationship of the group and the evolution of key morphological characters.

Cleaning old cabinets: revealing the taxonomy of Sri Lankan wolf spiders (Araneae, Udubidae and Zoropsidae).

The fauna of Udubidae and Zoropsidae of Sri Lanka is reviewed, the families are diagnosed and a distribution map is provided. The udubid genus Campostichomma Karsch, 1892 is diagnosed and redescribed, C. manicatum Karsch, 1892 is redescribed, and we propose three new species, based on females: C. harasbedda sp. nov., C. mudduk sp. nov. and C. alawala sp. nov. The zoropsid genus Devendra Lehtinen, 1967 is diagnosed and redescribed, D. seriatus (Simon, 1898), D. pumilus (Simon, 1898) and D. pardalis (Simon, 1898), are redescribed, and two new species are proposed: D. saama sp. nov. and D. amaiti sp. nov.

Description of Califorctenus (Cteninae, Ctenidae, Araneae), a new spider genus from Mexico.

Califorctenus gen. nov. (Araneae: Ctenidae) a new genus and a new species (Califorctenus cacachilensis sp. nov.) from the Baja California Peninsula in Mexico are described and illustrated. Califorctenus gen. nov. can be diagnosed by the morphology of the male palp, with embolus cylindrical and elongated, locking lobes of embolus positioned retrolaterally and the cup-shaped and elongated median apophysis.

Description of a new species of the Afrotropical spider genus Afroneutria (Araneae, Ctenidae).

A species of the Afrotropical spider genus Afroneutria is newly described. Afroneutria tanga sp. nov is endemic to Tanzania, southeast Africa, and can be distinguished from the remaining species of the genus by the retrolateral excavation of the embolus. An identification key for all Afroneutria species is presented.