taxonID	type	description	language	source
B45E4D47FFB4FFD2FC7CFA513798F8CF.taxon	discussion	Remarks: As discussed by Bond et al. (2012), the results of their molecular analyses did not recover a monophyletic Nemesiidae. This may be a product of insufficient taxon sampling, but more likely that the family is paraphyletic, as evidenced by Wheeler et al. (2017). Our dataset was not designed to test the hypothesis of nemesiid monophyly so we refrain from any further comments. TRIBE ANAMINI SIMON, 1889 Diagnosis: Members of the tribe Anamini differ from other nemesiids by the well-defined posterior heel on the maxillae (Figs 6 E, 7 E, 8 E, 9 E, 10 E, 11 E, 12 E, 13 E, 14 E, 15 E, 16 E, 17 E), the absence of claw tufts and a serrula and both males and females with two rows of teeth on the paired claws. Remarks: Our analyses (Fig. 3) confirm that species currently included in the genera Aname, Chenistonia, Hesperonatalius, Kwonkan, Merredinia, Namea, Pseudoteyl, Swolnpes, Teyl, Teyloides and Yilgarnia form a monophyletic group. This clade is defined by the presence of a well-defined posterior heel on the maxillae and equates to the Anamini as originally defined by Raven (1985 b). Other taxa that have been previously aligned with this group in the subfamily Anaminae, such as Stanwellia from Australasia, Acanthogonatus and Longistylus Indicatti & Lucas, 2005 from South America, Sinopesa Raven & Schwendinger, 1995 from Asia and Entypesa, Hermacha and Lepthercus from southern Africa and Madgascar, lack a pronounced maxillary heel (Raven, 1985 b; Raven & Schwendinger, 1995; Indicatti & Lucas, 2005; Indicatti et al., 2015). Stanwellia, Acanthogonatus and Longistylus, along with many other nemesiid taxa, possess pseudosegmented [or ‘ flexuous’ using the terminology of Goloboff (1995)] tarsi in males and some females (e. g. Raven, 1985 b; Goloboff, 1995; Indicatti & Lucas, 2005) and appear to be only distantly related to Anamini, as here defined (Bond et al., 2012; Wheeler et al., 2017). The classification proposed by Main (1985 a) recognized the tribes Anamini and Teylini. The Teylini included Teyl, Namea, Pseudoteyl and Teyloides and was defined by the embolus arising from the lateral side of the male pedipalpal bulb. In Anamini as so defined, the embolus arose from the distal end of the bulb. The lateral embolic configuration also occurs in Longistylus from South America (Indicatti & Lucas, 2005), but as Longistylus lacks the pronounced maxillary heel, it seems reasonable to assume that this represents a convergent morphology. Our analyses recovered a monophyletic group that includes Teyl, Namea and Pseudoteyl, as well as Merredinia, but the remaining Anamini were not recovered as reciprocally monophyletic (Fig. 3). Accordingly, we abandon the multiple-tribe classification and instead recognize several informal groups within Anamini: the Teyl group, the Kwonkan group, the Chenistonia group and the Aname group. We regard Teylini Main, 1982 as a junior synonym of Anamini Simon, 1899 (syn. nov.).	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFBAFFD3FE92FF7433C4FE41.taxon	discussion	Remarks: The analyses recovered a monophyletic Chenistonia group (Fig. 3) that consists of two reciprocally monophyletic genera, Chenistonia and Proshermacha, and a third lineage consisting of the monotypic genus Teyloides.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFBAFFD0FEC5FE76308FF9A2.taxon	description	FIG. 6 Chenistonia Hogg, 1901: 262. Type species: Chenistonia maculata Hogg, 1901, by original designation. Diagnosis: Males of Chenistonia s. s. differ from all other Anamini by the pear-shaped bulb with a short embolus (Fig. 6 C, D). Chenistonia further differs as follows: from Aname and Hesperonatalius by the maxillary cuspules being restricted to a relatively narrow band near the mesal edge (Fig. 6 B); from Aname by the lack of a ventral, asetose depression on the male pedipalpal tibia (Fig. 6 D); from Swolnpes by the digitiform tarsus I in males (Fig. 6 F); from Kwonkan and Swolnpes by the lack of thickened setae on the retrolateral face of the male pedipalpal tibia (Fig. 6 C) and lack of accessory branches on the spermathecae (Fig. 6 G); and from most Teyl, Teyloides and most Namea by the embolus arising from the distal end of the pedipalpal bulb (Fig. 6 C, D) or lacking a bulbous projection on metatarsus I (Fig. 6 F). Description: Small to medium nemesiid spiders. Coloration: sclerotized regions yellow-brown to brown, abdomen mottled. Cephalothorax: Carapace (Fig. 6 A) sparsely hirsute, with eight eyes in two rows; posterior median eyes (PME) slightly smaller than other eyes; fovea straight. Maxilla (Fig. 6 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, not restricted to narrow band; maxillary serrula absent. Labium (Fig. 6 E) wider than long, slightly indented anteriorly, without cuspules. Coxal cuspules absent (Fig. 6 B). Sternum (Fig. 6 B) with three pairs of small sigilla, close to lateral margin. Chelicera: Rastellum absent; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence absent. Pedipalp (Fig. 6 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) long and slender, with pronounced medial constriction (in lateral view); with pear-shaped bulb and very short embolus; embolus not reflexed. Legs: Male tibia I (Fig. 6 F) with large ventral spur bearing one or occasionally two megaspines; metatarsus I weakly incrassate; scopula usually present on entire ventral tarsi of legs I and II, and lighter scopula on tarsi III and IV, and metatarsi I and II; tarsi without spines; tarsus I not inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Fig. 6 G): One pair of low, rounded spermathecae. Distribution: Species of Chenistonia are known from the temperate mesic zone of south-eastern Australia (including Tasmania), tropical north-eastern Queensland and from the far south-western corner of Western Australia. Remarks: Our analyses demonstrate a significant genetic division between taxa previously thought to represent Chenistonia s. l. One group has males with a short embolus and a pear-shaped bulb (Fig. 6 C) and is consistent with the Aname maculata group as defined by Raven (1984 b). The other has a long embolus and rounded bulb (Fig. 7 C) and fits the Chenistonia tepperi ‘ super-species’ group defined by Main (1982 a). Due to differences in pedipalpal morphology and consistent, substantial genetic divergences (Fig. 3), we propose that these groups be recognized as separate genera, the first taking the name Chenistonia s. s. and the second taking the name Proshermacha, which is here removed from the synonymy of Chenistonia. The status and limits of Chenistonia s. l. has vacillated in the past, with the genus being treated as a junior synonym of Aname by Raven (1981, 1984 a, 1985 a) or as a distinct genus by Main (1982 a, 1985 b, 2012) and Raven (2000). The systematic position of the species here attributed to Chenistonia s. s. is confirmed due to the good descriptions provided by Raven (1984 b) and Main (2012). Included species: Chenistonia boranup Main, 2012; C. caeruleomontana (Raven, 1984); C. earthwatchorum (Raven, 1984); C. hickmani (Raven, 1984); C. maculate Hogg, 1901; C. montana (Raven, 1984); C. trevallynia Hickman, 1926; and C. tropica (Raven, 1984).	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFB9FFD1FF7CF905364FF8E1.taxon	description	FIG. 7	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFB9FFD1FF7CF905364FF8E1.taxon	diagnosis	Diagnosis: Species of Proshermacha differ from Chenistonia s. s. by the long embolus in males (Fig. 7 C, D). They also differ from other Anamini as follows: from Aname and Hesperonatalius by the maxillary cuspules being restricted to a relatively narrow band near the mesal edge (Fig. 7 B); from Kwonkan and Swolnpes by the lack of thickened setae on the retrolateral face of the male pedipalpal tibia (Fig. 7 C) and the lack of an accessory receptaculum in the female spermathecae (Fig. 7 G); from Swolnpes by the digitiform tarsus I in males (Fig. 7 F); and from Teyl, Teyloides and Namea by the embolus arising from the distal end of the pedipalpal bulb (Fig. 7 C) or lacking a bulbous projection on metatarsus I (Fig. 7 F). Description: Medium to large nemesiid spiders. Coloration: ranging from pale to dark brown. Cephalothorax: Carapace (Fig. 7 A) strongly hirsute, with eight eyes in two rows; PME slightly smaller than other eyes; fovea straight to procurved. Maxilla (Fig. 7 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, restricted to a narrow medial band; maxillary serrula absent. Labium (Fig. 7 E) wider than long, slightly indented anteriorly, without cuspules. Coxal cuspules absent (Fig. 7 B). Sternum (Fig. 7 B) with three pairs of sigilla; posterior pairs oval, subcentral. Chelicera: Rastellum weak or absent; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence absent. Pedipalp (Fig. 7 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) long and slender, with prominent medial constriction (in lateral view); with simple pyriform bulb and tapering embolus; embolus not reflexed. Legs: Male tibia I (Fig. 7 F) with large ventral spur bearing one, or occasionally two, megaspines; metatarsus I strongly incrassate; scopula usually present on entire ventral tarsi of legs I and II, and lighter scopula on tarsi III and IV, and metatarsi I and II; tarsi without spines; tarsus I not inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Fig. 7 G): One pair of spermathecae of variable shape. Distribution: Species of Proshermacha are known from southern South Australia and south-western Western Australia, in mesic, semi-arid and arid habitats. Remarks: As discussed above under Chenistonia, morphological and molecular data provide sufficient evidence for the resurrection of Proshermacha, which has long been considered a junior synonym of Aname or Chenistonia (Raven, 1981, 1984 a, 1985 a, 2000; Main, 1982 a, 1985 b, 2012). The consistent differences in the length of the embolus support each clade morphologically. Many of the previously named species here attributed to Proshermacha are poorly described and the status of those described by Hogg (1902), Simon (1908), Rainbow & Pulleine (1918) and Main (1954), and included in Chenistonia by Main (1982 a, 1985 b, 2012), are provisional and await confirmation based on revisionary studies. Indeed, the precise identity of the type species Proshermacha subarmata Simon, 1908, from south-western Australia, may remain unknown until additional adult specimens from the type locality can be identified. For this reason, we have not provided images of the type species, and instead include images of another species of Proshermacha (Fig. 7). Preliminary results from an unpublished molecular analysis strongly indicate that Proshermacha has high species-level diversity in south-western Australia (Harvey & Huey, unpublished data), which is confirmed by morphological features, primarily in the male pedipalp and the morphology of the first leg. This diversity has also been recently acknowledged by Main (2012). As it is very unlikely that any of these Western Australian species are conspecific with C. tepperi, which was originally described from southern South Australia (Hogg, 1902), we here remove all species from the synonymy of C. tepperi until species-level revisions are undertaken. Included species: Proshermacha auropilosa (Rainbow & Pulleine, 1918) comb. nov., transferred from Chenistonia; P. cuspidata (Main, 1954) comb. nov., transferred from Chenistonia; P. intricata (Rainbow & Pulleine, 1918) comb. nov., transferred from Aname; P. maculata (Rainbow & Pulleine, 1918) comb. nov., transferred from Aname; P. subarmata Simon, 1908, removed from synonymy with Chenistonia tepperi; P. tepperi (Hogg, 1902) comb. nov., transferred from Chenistonia; P. tigrina Simon, 1908, removed from synonymy with Chenistonia tepperi; and P. villosa (Rainbow & Pulleine, 1918) comb. nov., transferred from Chenistonia.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFBFFFD7FEC5FF7430ABF97A.taxon	description	FIG. 8	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFBFFFD7FEC5FF7430ABF97A.taxon	diagnosis	Diagnosis: Teyloides bakeri differs from all other Anamini except species of Teyl and Namea by the embolus arising from the retrolateral side of the pedipalpal bulb (Fig. 8 C, D) – a morphology which is presumably homoplastic given the phylogenetic placement of Teyloides in the Chenistonia group rather than the Teyl group. It differs from Teyl and most species of Namea by the presence of a megaspur and an enlarged spine on the tibia I of males (Fig. 8 F), and by the coiled spermathecae in females (Fig. 8 G) (Main, 1985 a). It can be further distinguished from most species of Namea by the absence of a stout and thick or long spine on the retroventral edge of the male pedipalpal tibia (see Raven, 1984 a). Description: Large nemesiid spiders. Coloration: generally yellow-brown. Cephalothorax: Carapace (Fig. 8 A) sparsely hirsute, with eight eyes in two rows; PME slightly smaller than other eyes; fovea straight. Maxilla (Fig. 8 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, not restricted to narrow band; maxillary serrula absent. Labium (Fig. 8 E) wider than long, slightly indented anteriorly, without cuspules. Coxal cuspules absent (Fig. 8 B). Sternum (Fig. 8 B) with three pairs of sigilla; posterior pairs rounded; all marginal. Chelicera: Rastellum absent; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence absent. Pedipalp (Fig. 8 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) long and slender, with distinct medial constriction (in lateral view); with rounded bulb and long, tapering embolus; embolus reflexed, arising from retrolateral side of bulb. Legs: Male tibia I (Fig. 8 F) with large ventral spur bearing one, or occasionally two, megaspines; metatarsus I strongly incrassate; scopula usually present on entire ventral tarsi of legs I and II, and lighter scopula on tarsi III and IV, and metatarsi I and II; tarsi without spines; tarsus I not inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Distribution: Teyloides bakeri is currently known only from southern South Australia, in the mesic Mount Lofty Range east of Adelaide. Remarks: The specimen of Teyloides bakeri in our molecular analyses was found to group with the Chenistonia group (Fig. 3). This was an initially unexpected result given the Teyl - like morphology of the male pedipalpal bulb, however, molecular data and the presence of a large leg I megaspur provide strong evidence of its affinities to the Chenistonia group.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFBEFFD7FBDDFBD63678FA64.taxon	discussion	Remarks: The Teyl group includes species previously included in Namea, Merredinia, Pseudoteyl and Teyl (Fig. 3). All except Merredinia have been previously considered members of the Teylini (Main, 1985 a) as they possess an embolus that arises from the lateral side of the pedipalpal bulb (Figs 9 C, 11 C, 12 C); a morphology homoplastic in the unrelated genus Teyloides. As discussed below, various species of Merredinia also fall within this group concept.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFBEFFEAFC0EF9B63082FAA8.taxon	description	FIGS 9 – 11	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FFBEFFEAFC0EF9B63082FAA8.taxon	diagnosis	Diagnosis: Most species of Teyl, Namea and Teyloides have the embolus arising from the lateral (usually retrolateral) side of the pedipalpal bulb (Figs 9 C, 11 C, 12 C), whereas in other Anamini it arises distally. The sole species of Teyl without this feature (Fig. 10 C), T. damsonoides (here transferred from Merredinia), has a sinuate metatarsus I which also has a rounded process (Fig. 10 F). Teyl species differ from Teyloides by the lack of a megaspur on the tibia I of males and the lack of coiled spermathecae in females, and from most species of Namea by the absence of a stout and thick or long spine on the retroventral edge of the male pedipalpal tibia (see Raven, 1984 a). Description: Small to large nemesiid spiders. Coloration: ranging from pale to dark brown. Cephalothorax: Carapace (Figs 9 A, 10 A, 11 A) strongly to sparsely hirsute, with eight eyes in two rows; PME slightly smaller than other eyes; fovea straight to slightly recurved. Maxilla (Figs 9 E, 10 E, 11 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, not restricted to narrow band; maxillary serrula absent. Labium (Figs 9 E, 10 E, 11 E) wider than long, slightly indented anteriorly, without cuspules. Coxal cuspules absent (Figs 9 B, 10 B, 11 B). Sternum (Figs 9 B, 10 B, 11 B) with three pairs of small sigilla; posterior pairs round to oval, marginal to subcentral. Chelicera: Rastellum weak to absent; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence soft and pallid. Pedipalp (Figs 9 C, D, 10 C, D, 11 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) long and slender, with medial constriction (in lateral view); with simple pyriform bulb and tapering embolus; embolus usually reflexed, but occasionally not reflexed. Legs: Male tibia I (Figs 9 F, 10 F, 11 F) without ventral spur, with one or more larger ventral spines; metatarsus I sometimes weakly incrassate, or sinuate; scopula usually present on entire ventral tarsi of legs I and II, and lighter scopula on tarsi III and IV, and metatarsi I and II; tarsi without spines; tarsus I not inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Figs 9 G, 10 G, 11 G): One pair of spermathecae of variable shape. Distribution: Species of Teyl are known from throughout southern and central Australia, in both mesic and arid habitats. Along with Aname and Kwonkan, they are a major component of the mygalomorph fauna of the central inland arid zone. Remarks: The molecular analyses recovered a clade containing species of Teyl, Pseudoteyl and Merredinia that was sister to Namea flavomaculata (Rainbow & Pulleine, 1918) (Fig. 3). This group did not include reciprocally monophyletic clades for all three named genera, and we therefore treat Pseudoteyl and Merredinia as junior synonyms of Teyl (syn. nov.). This decision is supported by the morphological data: the original diagnoses of Teyl and Pseudoteyl suggest only slight morphological disparities (Main, 1985 a), which we here suggest are insufficient to maintain separate genera. The morphological differences between Teyl and Merredinia are, however, more substantial. The embolus of Merredinia damsonoides (the sole named species of the genus) arises from the distal end of the pedipalpal bulb (Fig. 10 C, D), and the leg I of males has a sinuate metatarsus which also bears a bare rounded process (Fig. 10 F). More remarkably, an unusual male of Teyl MYG 455 from near Ravensthorpe (WAM T 72719) was included in a clade with the Merredinia specimens (Fig. 3) and has features of both genera, that is, it has metatarsus I with a bare rounded knob (which fits the diagnosis of Merredinia), but has a male pedipalp with the embolus arising from a lateral position (which matches the diagnosis of Teyl). The molecular data suggest that species of Merredinia have been derived from a Teyl - like ancestor, and that the male embolus has moved from a lateral position on the bulb to a terminal one, as in the taxa traditionally attributed to the Anamini. Indeed, Teyl MYG 455 resembles T. yeni Main, 2004 from western Victoria, which also has a curved metatarsus I with a rounded knob and an embolus arising from the retrolateral side of the bulb (Main, 2004). These data confirm that Merredinia is a junior synonym of Teyl (syn. nov.), a result which is further consistent with the observations of Main (1985 a, 2004), who suggested that Merredinia may indeed be misplaced in the Anamini due to the lack of a large spur on the tibia I of males. Included species: Teyl damsonoides (Main, 1983) comb. nov., transferred from Merredinia; T. harveyi Main, 2004; T. luculentus Main, 1975; T. vancouveri (Main, 1985 a) comb. nov., transferred from Pseudoteyl; T. walkeri Main, 2004; and T. yeni Main, 2004.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF83FFEBFEF2FA1A3368F943.taxon	description	FIG. 12	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF83FFEBFEF2FA1A3368F943.taxon	diagnosis	Diagnosis: Species of Namea closely resemble Teyloides and Teyl in that males of most species have the embolus arising from the lateral side of the pedipalpal bulb (Fig. 12 C, D). Namea can be distinguished from Teyloides by the absence in most species of a megaspur with an enlarged spine on the tibia I of males, and by the lack of coiled spermathecae in females. Namea can be further distinguished from species of Teyl and Teyloides by the presence in all but N. dahmsi Raven, 1984 and N. excavans Raven, 1984 of a stout and thick or long spine on the retroventral edge of the male pedipalpal tibia. Description: Small to medium-sized nemesiid spiders. Coloration: usually yellow-brown to brown. Cephalothorax: Carapace (Fig. 12 A) usually hirsute, with eight eyes in two rows; PME usually slightly smaller than other eyes; fovea straight to slightly procurved. Maxilla (Fig. 12 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, not restricted to narrow band; maxillary serrula absent. Labium (Fig. 12 E) wider than long, slightly indented anteriorly, usually without cuspules. Coxal cuspules absent (Fig. 12 B). Sternum (Fig. 12 B) with three pairs of sigilla; posterior pairs usually rounded to oval, submarginal. Chelicera: Rastellum sometime present; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence soft and pallid. Pedipalp (Fig. 12 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) long, with distinct medial constriction (in lateral view); with simple pyriform bulb and tapering embolus; embolus not reflexed. Legs: Male tibia I (Fig. 12 F) usually without large ventral spur, with one or more megaspines; metatarsus I with slight proximal excavation; scopula usually present on entire ventral tarsi of legs I and II, and lighter scopula on tarsi III and IV, and metatarsi I and II; tarsi without spines; tarsus I not inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Fig. 12 G): One pair of spermathecae of variable shape. Distribution: Species of Namea are known from the mesic forests of eastern Australia, with a range that extends from the Wet Tropics of north-eastern Queensland south to mid-eastern New South Wales (Raven, 1984 a). Remarks: When first described, Namea was hypothesized to be most similar to Teyl (Raven, 1984 a). It was formally included in the tribe Teylini by Main (1985 a), based on the embolus originating on the retrolateral side of the bulb in the majority of species. The only species where the embolus arose distally, N. callemonda Raven, 1984 and N. dahmsi from south-eastern Queensland, were only doubtfully referred to Namea by Raven (1984 a) and Main (1985 a). The single exemplar of Namea in our analysis, N. flavomaculata, was recovered as the sister-group to the genus Teyl (Fig. 3). Further taxa, especially the morphologically anomalous N. callemonda and N. dahmsi, are required to test the limits and monophyly of this diverse genus. Included species: Namea brisbanensis Raven, 1984; N. bunya Raven, 1984; N. calcaria Raven, 1984; N. callemonda Raven, 1984; N. capricornia Raven, 1984; N. cucurbita Raven, 1984; N. dahmsi Raven, 1984; N. dicalcaria Raven, 1984; N. excavans Raven, 1984; N. flavomaculata (Rainbow & Pulleine, 1918); N. jimna Raven, 1984; N. Nebulosa Raven, 1984; N. olympus Raven, 1984; N. salanitri Raven, 1984; and N. saundersi Raven, 1984.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF82FFEBFE89F96836ADFD55.taxon	discussion	Remarks: The analysis recovered a monophyletic group of species including the genera Kwonkan, Yilgarnia and Swolnpes, as well as Aname turrigera Main, 1994 (Fig. 3). Females of this group possess a small and apparently synapomorphic medial accessory receptaculum on the spermathecae (Main, 1983; Raven, 1984 b; Castalanelli et al., 2014) (Figs 13 G, 14 G, 15 G), and males have a distinctive field of spinules on the retrolateral face of the pedipalpal tibia (Main, 1983, 1986, 2008; Main & Framenau, 2009) (Figs 13 C, 14 C, 15 C). Whether these modified setae also occur in A. turrigera will depend on males being identified, as it was originally described only from females (Main, 1994). Our analysis found good support for the Kwonkan group, but the internal relationships were less well resolved (Fig. 3). Species of Swolnpes were sister to the other taxa (Fig. 3), but Kwonkan and Yilgarnia are treated as synonyms (see below).	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF82FFE9FC12FD0C36A7FABA.taxon	description	FIGS 13, 14	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF82FFE9FC12FD0C36A7FABA.taxon	diagnosis	Diagnosis: Species of Kwonkan differ from other Anamini except Swolnpes by the presence of a field of spinules on the retrolateral face of the male pedipalpal tibia (Figs 13 C, 14 C), and a small accessory receptaculum on the female spermathecae (Figs 13 G, 14 G). Kwonkan differs from Swolnpes by the digitiform tarsus I in males (Figs 13 F, 14 F). Description: Small to medium nemesiid spiders. Coloration: ranging from pale to yellow-brown. Cephalothorax: Carapace (Figs 13 A, 14 A) strongly to sparsely hirsute, with eight eyes in two rows; PME slightly smaller than other eyes; fovea straight to procurved. Maxilla (Figs 13 E, 14 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, not restricted to narrow band; maxillary serrula absent. Labium (Figs 13 E, 14 E) wider than long, slightly indented anteriorly, without cuspules. Coxae III and IV usually without cuspules, but sometimes present. Sternum (Figs 13 B, 14 B) with three pairs of sigilla; posterior pairs oval to elongate, marginal to submarginal. Chelicera: Rastellum weak to stout; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence absent. Pedipalp (Figs 13 C, D, 14 C, D): Male tibia uniformly setose, with patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) short and terminally blunt, without medial constriction (in lateral view); with simple pyriform bulb and tapering embolus; embolus not reflexed. Legs: Male tibia I (Figs 13 F, 14 F) with large ventral spur bearing one, or occasionally two, megaspines; metatarsus I strongly incrassate; scopula usually present on entire ventral tarsi of legs I and II, and lighter scopula on tarsi III and IV, and metatarsi I and II; tarsi usually with one or more spines; tarsus I not inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Figs 13 G, 14 G): One pair of lobate spermathecae each with small accessory receptaculum. Distribution: Species of Kwonkan are known from throughout southern and central Australia, mostly in arid or semi-arid habitats. Along with Aname and Teyl, they are a major component of the mygalomorph fauna of the central inland arid zone. Remarks: The molecular analyses confirmed that Kwonkan and Yilgarnia formed a monophyletic group which also included Aname turrigera Main, 1994 (Fig. 3). The clade Kwonkan + Yilgarnia was sister to Swolnpes. Females of all three taxa possess a small, medial accessory receptaculum on the spermathecae (Main, 1983, 1986, 1994) (Figs 13 G, 14 G, 15 G), and males have a field of spinules on the retrolateral face of the pedipalpal tibia (Main, 1983, 1986, 2008) (Figs 13 C, 14 C, 15 C). While Kwonkan and Yilgarnia were originally defined on putatively autapomorphic features including the presence of spines on at least one pair of pedal tarsi in Kwonkan (Main, 1983) and cuspules on coxae III and IV in Yilgarnia (Main, 1986, 2008) (Fig. 14 B), we have examined specimens (lodged in the WAM) that confound these diagnoses, including some species that lack tarsal spines and others that have both tarsal spines and coxal cuspules. Rather than devise a generic classification that attempts to incorporate all of this variation, we prefer to recognize a single genus-group name, Kwonkan, and redefine the genus as outlined in the Diagnosis (above). This renders Yilgarnia as a junior synonym of Kwonkan (syn. nov.). The molecular inclusion of Aname turrigera in Kwonkan is also strongly supported by the morphological features of the females described by Main (1994). The spermathecae have a small accessory receptaculum as found in other species of the Kwonkan group (Main, 1994, fig. 1 I – K), and the abdomen has a distinctive colour pattern with a series of dark chevrons on a pale background, reminiscent of patterns found in other congeners (Main, 1983, 1986, 2008). Main (1994) noted a strong resemblance between A. turrigera, Kwonkan and Yilgarnia, but as A. turrigera lacked the diagnostic features of those genera, which included the presence of tarsal spines in Kwonkan and coxal cuspules in Yilgarnia, she suggested it may eventually be found to belong to a new genus. Aname turrigera has been recorded from either side of the Nullarbor Plain of southern Australia and is the only nemesiid that is known to construct an elevated turret to its burrow through or against vegetation (Main, 1994) (Fig. 1 O). The genus Kwonkan now contains nine named species including the six species originally included in the genus Kwonkan by Main (1983), two species formerly in Yilgarnia (Main, 1986, 2008) and K. turrigera (Main, 1994). There are numerous other species known from museum collections, and there is ample molecular data to support the hypothesis of a highly diverse fauna (Castalanelli et al., 2014; this study). Included species: Kwonkan anatolion Main, 1983; K. currycomboides (Main, 1986) comb. nov., transferred from Yilgarnia; K. eboracum Main, 1983; K. goongarriensis Main, 1983; K. linnaei (Main, 2008) comb. nov., transferred from Yilgarnia; K. moriartii Main, 1983; K. silvestris Main, 1983; K. turrigera (Main, 1994) comb. nov., transferred from Aname; and K. wonganensis (Main, 1977).	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF80FFEEFCBEFA1F376DF8E0.taxon	description	FIG. 15	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF80FFEEFCBEFA1F376DF8E0.taxon	diagnosis	Diagnosis: Species of Swolnpes differ from all other Anamini by the inflated tarsus I in males (Fig. 15 F). Description: Small nemesiid spiders. Coloration: mostly pale yellow. Cephalothorax: Carapace (Fig. 15 A) weakly hirsute, with eight eyes in two rows; PME slightly smaller than other eyes; fovea slightly procurved. Maxilla (Fig. 15 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, not restricted to narrow band; maxillary serrula absent. Labium (Fig. 15 E) wider than long, slightly indented anteriorly, without cuspules. Coxal cuspules absent (Fig. 15 B). Sternum (Fig. 15 B) with three pairs of barely discernible sigilla; posterior pairs elongate, marginal. Chelicera: absent; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence absent. Pedipalp (Fig. 15 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) short and terminally blunt, without medial constriction (in lateral view); with simple pyriform bulb and tapering embolus; embolus not reflexed. Legs: Male tibia I (Fig. 15 F) without ventral spur; metatarsus I not incrassate; scopula usually present on entire ventral tarsi of legs I and II, and lighter scopula on tarsi III and IV, and metatarsi I and II; tarsi without spines; tarsus I inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Fig. 15 G): One pair of small lobate spermathecae with accessory receptaculum. Distribution: Species of Swolnpes are known only from the arid zone of southern Western Australia (see Main & Framenau, 2009). Remarks: As noted above, the molecular analyses show that the genus Swolnpes is the sister group to Kwonkan. The two named species of Swolnpes and at least two undescribed species (S. MYG 234 and S. MYG 415) are known from museum collections (Main & Framenau, 2009; this study). Included species: Swolnpes darwini Main & Framenau, 2009; and S. morganensis Main & Framenau, 2009.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF86FFEFFEB9FBF832E1FA35.taxon	discussion	Remarks: Our analysis recovered a strongly monophyletic Aname group that was sister to the Kwonkan group (Fig. 3). The group was in turn divided into two main clades, the first representing the genus Aname containing males with a megaspine and an enlarged spur on leg I, and Hesperonatalius for males that lack these features. Despite the close morphological similarity of Aname with species formerly included in Chenistonia (but here referred to Proshermacha), we found that Aname never grouped with Proshermacha in the molecular analyses (Fig. 3).	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF86FFE2FEFAF9BA3331FEC5.taxon	description	FIG. 16	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF86FFE2FEFAF9BA3331FEC5.taxon	diagnosis	Diagnosis: Species of Aname differ from all other Australasian anamine genera by the presence of a ventral asetose depression on the male pedipalpal tibia (Fig. 16 D). It further differs from some of these genera as follows: from Kwonkan and Swolnpes by the absence of a field of spinules on the retrolateral face of the male pedipalpal tibia (Fig. 16 C) and the lack of an accessory receptaculum on the spermathecae (Fig. 16 G); from Swolnpes by the digitiform tarsus I in males (Fig. 16 F); and from members of the Teyl group by the embolus arising from the distal end of the bulb (Fig. 16 C) or by the metatarsus I of males being straight and without a rounded process (Fig. 16 F). Description: Small to large nemesiid spiders. Coloration: ranging from pale to dark brown. Cephalothorax: Carapace (Fig. 16 A) strongly to sparsely hirsute, with eight eyes in two rows; PME slightly smaller than other eyes; fovea straight to procurved. Maxilla (Fig. 16 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, not restricted to narrow band; maxillary serrula absent. Labium (Fig. 16 E) wider than long, slightly indented anteriorly, without cuspules. Coxal cuspules absent (Fig. 16 B). Sternum (Fig. 16 B) with three pairs of sigilla; posterior pairs oval to elongate, marginal to subcentral. Chelicera: Rastellum weak to absent; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence absent. Pedipalp (Fig. 16 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face (but larger spines sometimes present), and without asetose ventral depression; tarsus (cymbium) short and terminally blunt, without medial constriction (in lateral view); with simple pyriform bulb and tapering embolus; embolus not reflexed. Legs: Male tibia I (Fig. 16 F) with large ventral spur bearing one, or occasionally two, megaspines; metatarsus I strongly incrassate; scopula usually present on entire ventral tarsi of legs I and II, and lighter scopula on tarsi III and IV, and metatarsi I and II; tarsi without spines; tarsus I not inflated; usually three claws on all legs, sometimes two on legs I, II, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth, absent on leg I of some species. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Fig. 16 G): One pair of spermathecae of variable shape. Distribution: Species of Aname occur throughout most of Australia, including Tasmania, in a wide array of both arid and mesic habitats. Along with Kwonkan and Teyl, they are a major component of the mygalomorph fauna of the central inland arid zone. Remarks: The molecular analyses recovered a monophyletic assemblage of Aname species, sister to Hesperonatalius (Fig. 3). Aname has three junior synonyms: Dekana Hogg, 1902, Dolichosternum Rainbow & Pulleine, 1918 and Sungenia Rainbow & Pulleine, 1918. All three taxa were synonymized by Raven (1981), along with Chenistonia, although the latter has since been treated as a valid genus (e. g. Main, 1982 a, 1985 b, 2012; Raven, 2000; this study). Although we were unable to include specimens of the type species of these synonymous genera in our analysis, we are confident that their synonymy with Aname is warranted based on the morphological criteria discussed by Raven (1981). As noted elsewhere, our data firmly demonstrate that Chenistonia is quite distinct from Aname and is confirmed as a valid genus (see above). The genus Aname occurs widely across mainland Australia andTasmania, and includes 30 named species, as well as numerous unnamed species (e. g. Castalanelli et al., 2014; this study). Along with Idiosoma, it may be among the most diverse of the Australian mygalomorph genera (Huey, Rix & Harvey, unpublished data). Included species: Aname aragog Harvey, Framenau, Wojcieszkek, Rix & Harvey, 2012; A. armigera Rainbow & Pulleine, 1918; A. atra (Strand, 1913); A. aurea Rainbow & Pulleine, 1918; A. barrema Raven, 1985; A. blackdownensis Raven, 1985; A. camara Raven, 1985; A. carina Raven, 1985; A. coenosa Rainbow & Pulleine, 1918; A. collinsorum Raven, 1985; A. comosa Rainbow & Pulleine, 1918; A. distincta (Rainbow, 1914) (= Dolichosternum attenuatum Rainbow and Pulleine; 1918, A. villosa Rainbow & Pulleine, 1918); A. diversicolor (Hogg, 1902); A. ellenae Harvey, Framenau, Wojcieszkek, Rix & Harvey, 2012; A. fuscocincta Rainbow & Pulleine, 1918; A. grandis Rainbow & Pulleine, 1918 (= A. robusta Rainbow and Pulleine; 1918); A. hirsuta Rainbow & Pulleine, 1918; A. humptydoo Raven, 1985; A. inimica Raven, 1985; A. kirrama Raven, 1984; A. longitheca Raven, 1985; A. maculata Rainbow & Pulleine, 1918; A. mainae Raven, 2000; A. marae Harvey, Framenau, Wojcieszkek, Rix & Harvey, 2012; A. mellosa Harvey, Framenau, Wojcieszkek, Rix & Harvey, 2012; A. pallida L. Koch, 1873 (= Chenistonia giraulti Rainbow, 1914); A. platypus (L. Koch, 1875); A. robertsorum Raven, 1985; A. tasmanica Hogg, 1902; A. tigrina Raven, 1985; A. villosa (Rainbow & Pulleine, 1918); and A. warialda Raven, 1985.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF86FFE2FEFAF9BA3331FEC5.taxon	diagnosis	Diagnosis: Species of Hesperonatalius differ from all other Australasian anamine genera as follows: from Kwonkan and Swolnpes by the lack of spinules on the retrolateral face of the male pedipalpal tibia (Fig. 17 C) (and presumably the lack of an accessory receptaculum on the spermathecae, but females are currently unknown); from Swolnpes by the digitiform tarsus I in males (Fig. 17 F); from most species of the Teyl group by the embolus arising from the distal end of the bulb (Fig. 17 C, D), or the male first leg being straight and without a rounded knob (Fig. 17 F); from Chenistonia, Proshermacha and Aname by the lack of a megaspur and enlarged spine on the tibia I of males (Fig. 17 F); and from Aname by the lack of a ventral asetose depression on the male pedipalpal tibia (Fig. 17 D). Description: Small nemesiid spiders. Coloration: pale to yellow-brown. Cephalothorax: Carapace (Fig. 17 A) sparsely hirsute, with eight eyes in two rows; PME slightly smaller than other eyes; fovea straight. Maxilla (Fig. 17 E) with strongly produced basal heel; with numerous cuspules distributed over medial half and heel of each maxilla, not restricted to narrow band; maxillary serrula absent. Labium (Fig. 17 E) wider than long, slightly indented anteriorly, without cuspules. Coxal cuspules absent (Fig. 17 B). Sternum (Fig. 17 B) either without sigilla or with one elongate posterior pair, marginal. Chelicera: Rastellum absent; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence absent. Pedipalp (Fig. 17 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) short and terminally blunt, without medial constriction (in lateral view); with simple pyriform bulb and tapering embolus; embolus not reflexed. Legs: Male tibia I (Fig. 17 F) with low ventral spur bearing one megaspine; metatarsus I weakly incrassate; very sparse scopula present on metatarsus and tarsi of legs; tarsi without spines; tarsus I not inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Fig. 17 G): Unknown. Distribution: Species of Hesperonatalius are known only from the arid western margin of Western Australia, with a range that extends from Lake MacLeod to Geraldton. Remarks: We recovered Hesperonatalius as the sister genus to Aname in our molecular phylogeny (Fig. 3), from which it primarily differs by the lack of a ventral asetose depression on the male pedipalpal tibia (Castalanelli et al., 2017) (Fig. 17 D). The genus currently contains three named species, but none are represented by females and the morphology of the spermathecae is unknown. I n c l u d e d s p e c i e s: H e s p e r o n a t a l i u s h a r r i e t a e C a s t a l a n e l l i, H u e y, H i l l y e r & H a r v e y, 2 0 1 7; H. langlandsi Castalanelli, Huey, Hillyer & Harvey, 2017; and H. maxwelli Castalanelli, Huey, Hillyer & Harvey, 2017.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF8BFFE3FF7BFEDB36AAF899.taxon	description	FIG. 18	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
B45E4D47FF8BFFE3FF7BFEDB36AAF899.taxon	diagnosis	Diagnosis: Stanwellia differs from other species of Australasian Anamini by the lack of a posterior heel on the maxilla, the absence of cuspules on the posteromesal corner of the maxilla (Fig. 18 B), and the presence of flexuous tarsi. D e s c r i p t i o n: S m a l l t o l a r g e n e m e s i i d s p i d e r s. Coloration: ranging from pale to dark brown. Cephalothorax: Carapace (Fig. 18 A) strongly to sparsely hirsute, with eight eyes in two rows; PME slightly smaller than other eyes; fovea straight to procurved. Maxilla without strongly produced basal heel; with numerous cuspules confined to anterior ectal corner; maxillary serrula usually present. Labium (Fig. 18 E) wider than long, slightly indented anteriorly, sometimes with cuspules. Coxal cuspules absent (Fig. 18 B). Sternum (Fig. 18 B) with three pairs of sigilla; posterior pairs rounded, situated near lateral margin. Chelicera: Rastellum usually present; cheliceral furrow with several prominent promarginal teeth and several small granules basomesally; intercheliceral tumescence absent. Pedipalp (Fig. 18 C, D): Male tibia uniformly setose, without patch of spinules on retrolateral face, and without asetose ventral depression; tarsus (cymbium) short and terminally blunt, without medial constriction (in lateral view); with simple pyriform bulb and tapering embolus; embolus not reflexed. Legs: Male tibia I (Fig. 18 F) without ventral spur; metatarsus I not incrassate; scopula usually present on all tarsi of males; tarsi flexuous and without spines; tarsus I not inflated; three claws, lateral claws each with two short rows of teeth; medial claw small and without ventral teeth. Abdomen: Longer than wide. Two pairs of spinnerets; posterior median spinnerets unsegmented and separated by about diameter of spinneret; posterior lateral spinnerets three-segmented, apical segment elongate, digitiform. Female genitalia (Fig. 18 G): One pair of spermathecae of lobate spermathecae. Remarks: Although considered a member of the Anaminae based on morphological criteria (Raven, 1985 b), alternative phylogenetic positions for Stanwellia have since been proposed. In a cladistic treatment based on morphological characters, Goloboff (1995) found Stanwellia to group with Acanthogonatus, Pycnothele, Rachias and Stenoterommata from South America and Hermacha from South Africa. This alternative placement for Stanwellia with Acanthogonatus and Stenoterommata was also recovered by Bond et al. (2012) and Leavitt et al. (2015). Bond et al. (2012) found these three genera to be sister to Fufius, a result that was also recovered in our analysis, albeit with weak support (Fig. 2). Wheeler et al. (2017) recovered a Stanwellia + Acanthogonatus clade as sister to all other Nemesiidae, while Leavitt et al. (2015) found the Stanwellia, Acanthogonatus, Stenoterommata clade to be sister to Pionothele, with Fufius sister to Brachythele Ausserer, 1871, from Europe. Our analyses consistently failed to recover Stanwellia as part of the Anamini (Fig. 3). In the three gene analysis (Fig. 2), it formed a group with Acanthogonatus and Stenoterommata, similar to the results of Bond et al. (2012) and Leavitt et al. (2015), and Raven (1985 b) also inferred a sister-group relationship between Stanwellia and Acanthogonatus. If a close relationship between Stanwellia, Acanthogonatus, Stenoterommata and Pycnothele is maintained, as proposed by Goloboff (1995), the subfamily-group name Pycnothelinae Chamberlin, 1917 is available for this clade (Chamberlin, 1917). The New Zealand specimens of Stanwellia used in this study nested within the Australian clade (Fig. 3), rendering Stanwellia as the only nemesiid genus that occurs in both Australia and New Zealand. The New Zealand species were previously included in the endemic genus Aparua (e. g. Todd, 1945; Forster & Wilton, 1968) until Main (1983) treated the latter as a junior synonym of Stanwellia. Included species: Australian species: Stanwellia annulipes (C. L. Koch, 1842); S. grisea (Hogg, 1901) (= Aname arborea Hogg, 1901; A. pellucida Hogg, 1901; I. gregori Hogg, 1901; Chenistonia major Hogg, 1901; A. butleri Rainbow & Pulleine, 1918); S. hoggi (Rainbow, 1914) (= S. decora Rainbow & Pulleine, 1918; Aname decora Rainbow & Pulleine, 1918); S. inornata Main, 1972; S. minor (Kulczynski, 1908); S. nebulosa (Rainbow & Pulleine, 1918) (= Aname confusa Rainbow & Pulleine, 1918); S. occidentalis Main, 1972; S. pexa (Hickman, 1930). New Zealand species: S. bipectinata (Todd, 1945); S. hapua (Forster, 1968); S. hollowayi (Forster, 1968); S. houhora (Forster, 1968); S. kaituna (Forster, 1968); S. media (Forster, 1968); S. puna (Forster, 1968); S. regia (Forster, 1968); S. taranga (Forster, 1968); S. tuna (Forster, 1968). PROBLEMATIC ANAMINE TAXA AND AVENUES FOR FUTURE RESEARCH While our taxon sampling of Australasian Anaminae (sensu Raven, 1985 b) is relatively broad, at least at the generic level, there are some taxa that require further research to test their systematic placement and answer lingering questions regarding their relationships. For example, the monophyly of the genus Namea could not be confirmed with our dataset, and the morphological features of some species preclude a straightforward diagnosis (Raven, 1984 a). In particular, two species from south-eastern Queensland, N. callemonda and N. dahmsi, lack the laterally originating embolus that occurs in other species of the genus (Raven, 1984 a) and in all other taxa assigned by Main (1985 a) to the Teylini. Raven (1984 a) and Main (1985 a) highlighted the anomalous morphology of both of these species and suggested that they might be better placed in a new genus. We recommend that sequence data be used to elucidate their relationships more effectively, and that a variety of Namea species will help to test the monophyly of this morphologically diverse assemblage of taxa. Similarly, females have not yet been attributed to any of the species of Hesperonatalius (Castalanelli et al., 2017), and males of Kwonkan turrigera are currently unknown, although our analysis hypothesizes that they will possess a field of spinules on the retrolateral face of the pedipalpal tibia, similar to other species of Kwonkan. Finally, the phylogenetic placement of the genus Stanwellia can now only be more fully resolved using multi-gene molecular datasets from a broad range of nemesiid taxa. While the analyses undertaken thus far suggests a close relationship with the South American genera Acanthogonatus and Stenoterommata, and possibly also Fufius and Pionothele (Bond et al., 2012; Leavitt et al., 2015), samples of many other taxa are required to adequately test these hypotheses. Such research will also help determine the monophyly and limits of the Nemesiidae, which may be at best paraphyletic, or at worst polyphyletic.	en	Harvey, Mark S, Hillyer, Mia J, Main, Barbara York, Moulds, Timothy A, Raven, Robert J, Rix, Michael G, Vink, Cor J, Huey, Joel A (2018): Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184 (2): 407-452, DOI: 10.1093/zoolinnean/zlx111, URL: https://academic.oup.com/zoolinnean/article/184/2/407/4932782
