Monograph |
Corresponding author: Xin Xu ( xuxin_09@163.com ) Academic editor: Jeremy Miller
© 2019 Xin Xu, Hirotsugu Ono, Matjaž Kuntner, Fengxiang Liu, Daiqin Li.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Xu X, Ono H, Kuntner M, Liu F, Li D (2019) A taxonomic monograph of the liphistiid spider genus Heptathela, endemic to Japanese islands. ZooKeys 888: 1-50. https://doi.org/10.3897/zookeys.888.34494
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Among the eight extant genera of primitively segmented spiders, family Liphistiidae, two are confined to East Asian islands, Heptathela Kishida, 1923 and Ryuthela Haupt, 1983. In this paper, a taxonomic revision of the genus Heptathela (Heptathelinae) from Kyushu and Ryukyu archipelago, Japan is provided. This study follows a multi-tier species delimitation strategy within an integrative taxonomic framework that is presented in a parallel paper, in which diagnosable lineages are considered as valid species. There, the initial hypothesis of species diversity (19) based on classical morphological diagnoses is tested with multiple species delimitation methods aimed at resolving conflict in data. This revision follows those analyses that converge on the species diversity of 20, which includes a pair of cryptic species that would have been undetected with morphology alone. After this revision, eight previously described species remain valid, two junior synonyms are proposed, and 12 new Heptathela species are described based on diagnostic evidence. To ease identification and to hint at putative evolutionary units, Heptathela is divided into three groups. The Kyushu group contains H. higoensis Haupt, 1983, H. kikuyai Ono, 1998, H. kimurai (Kishida, 1920), and H. yakushimaensis Ono, 1998; the Amami group contains H. amamiensis Haupt, 1983, H. kanenoi Ono, 1996, H. kojima sp. nov., H. sumiyo sp. nov., and H. uken sp. nov.; and the Okinawa group contains H. yanbaruensis Haupt, 1983, H. aha sp. nov., H. gayozan sp. nov., H. kubayama sp. nov., H. mae sp. nov., H. otoha sp. nov., H. shuri sp. nov., H. tokashiki sp. nov., H. unten sp. nov., and H. crypta sp. nov. Heptathela helios Tanikawa & Miyashita, 2014 is not assigned to a species group. A combination of diagnostic tools augments the morphological diagnoses that, in isolation, would be prone to error in morphologically challenging groups of organisms.
Heptathelinae, island endemism, Kyushu, Ryukyu archipelago, species delimitation, trapdoor spiders
The spider family Liphistiidae is the sole extant lineage within the ancient suborder Mesothelae known to possess a combination of plesiomorphic traits, for example, abdominal tergites (Fig.
Taxonomic limits of Heptathela were recently redefined (
Heptathela therefore does not occur in continental Asia (
We follow this evidence with a thorough taxonomic revision of Heptathela. As is the case in other liphistiids, Heptathela females usually lack clear morphologically diagnostic characters and furthermore exhibit considerable intraspecific variation. While males are more readily diagnosable morphologically, they are very rarely collected (
Our original sampling, performed in the entire Heptathela range from Kyushu to the central Ryukyu archipelago, relied on the type locality information of each known Heptathela species, but also involved previously unexplored areas, focusing particularly on roadside habitats (Figs
Microhabitats, trapdoors, and general somatic morphology of Heptathela Kishida, 1923 A–E Kyushu group: A microhabitat of H. kimurai (Kishida, 1920) at Shiroyama Park, Kyushu B, C open and closed trapdoor of H. kimurai D female H. kimurai E male H. higoensis (Haupt, 1983) F–J Amami group: F microhabitat of H. sumiyo sp. nov. at Sumiyo-cho, Amamioshima G, H open and closed trapdoor of H. amamiensis Haupt, 1983 I female H. sumiyo sp. nov. J male H. sumiyo sp. nov. K–O Okinawa group: K microhabitat of H. yanbaruensis Haupt, 1983 at Yona, Okinawajima L, M open and closed trapdoor of H. yanbaruensis N female H. yanbaruensis O male H. yanbaruensis.
Specimens were morphologically examined under an Olympus SZX16 stereomicroscope and an Olympus BX51 compound microscope. Their genitalia were cleared in boiling 10% KOH for a few minutes to dissolve soft tissues. Unless noted otherwise, left palps were imaged. All measurements are reported in millimetres. Leg and palp measurements are given in the following order: total length (femur + patella + tibia + metatarsus + tarsus). The value ranges for cheliceral groove denticles, spinnerets and measurements in taxonomic descriptions are based on all the examined specimens.
Taxonomic descriptions using morphological characteristics follow our established methodology (
ALE anterior lateral eyes,
AME anterior median eyes,
BL body length,
CL carapace length,
Co conductor,
CT contrategulum,
CW carapace width,
D depression,
E embolus,
OL opisthosoma length,
OW opisthosoma width,
PC paracymbium,
PLE posterior lateral eyes,
PME posterior median eyes,
RC receptacular cluster,
T tegulum.
Museum abbreviations:
CBEE Centre for Behavioural Ecology and Evolution, School of Life Sciences, Hubei University, Wuhan, China;
ZMH Zoological Museum Hamburg, Germany.
Voucher specimens are deposited at CBEE, and the type specimens will be deposited in
The integrative taxonomic framework considers diagnosable lineages as potentially valid species. Our taxonomic approach is based on morphological diagnoses that provide an initial species hypothesis (19 species). This is then further tested within a parallel, molecular species delimitation study (
The aligned DNA barcode gene COI is deposited in the Dryad Digital Repository.
Of the total species diversity of 20, eight previously described species remain valid, two junior synonyms are proposed, and 12 new Heptathela species are discovered and described. This taxonomic revision thus represents a 137.5% increase in species diversity of the genus, drawing parallels with prior species richness underestimation found also in other liphistiid genera, e.g., Ganthela (
To ease identification and to hint at putative evolutionary units, we divide Heptathela into the Kyushu, the Amami, and the Okinawa groups, each of which contains four, five, and ten species, respectively (Fig.
Type species.
Diagnosis. Heptathela males differ from Liphistius males by lacking the tibial apophysis, and from males of all other Heptathelinae genera by possessing a leaf-shaped conductor (Fig.
Male and female genital anatomy of Heptathela higoensis Haupt, 1983 A, C 3472 (short for XUX-2013-472) B, D 3435 E–G 3365 H–K 3381 A, B vulva dorsal view C, D vulva ventral view E palp prolateral view F palp ventral view G palp retrolateral view H–K palp distal view; 3365: Hitoyoshi Ruins Park, Kumamoto; 3472: Takachihokawara, Kagoshima; 3435: Mukoyama, Miyazaki; 3381: Kozomo, Kumamoto. Scale bar: 0.5 mm.
General description. All the 20 species share the following characteristics: sternum narrow, longer than wide; a few short pointed hairs running over ocular mound in a longitudinal row; chelicerae robust with promargin of cheliceral groove with variable sized denticles (the number of denticles is presented under each species description); legs with strong hairs and spines, and male legs distinctly longer than female legs; opisthosoma with 12 tergites, and the fifth tergite is the largest in all the species except in H. amamiensis, H. sumiyo sp. nov., and H. uken sp. nov., in which the fourth tergite is the largest.
Diagnosis. The males of the Kyushu group differ from those of the other two groups by the nearly rectangular contrategulum whose two proximal thirds are serrated (Fig.
Monophyly. The Bayesian analyses based on concatenated two genes and two partitions (for details, see
Composition. H. higoensis Haupt, 1983, H. kikuyai Ono, 1998, H. kimurai (Kishida, 1920), and H. yakushimaensis Ono, 1998.
Distribution. Kyushu and Yakishima (Fig.
Heptathela kimurai higoensis
Haupt, 1983: 283 (holotype: male, from Kumamoto, Kyushu, Japan, collected by M. Yoshikura on 27 September 1973, deposited in
Heptathela nishikawai
Ono, 1998: 19 (holotype: female, from Hitoyoshi-shi, Kumamoto-ken, Kyushu, Japan, collected by H. Ono on 19 November 1996, deposited in
Heptathela yaginumai
Ono, 1998: 20 (holotype: female, from Honjo, Kunitomi-cho, Higashimorokata-gun, Miyazaki-ken, Kyushu, Japan, collected by T. Yaginuma on 18 June 1949, deposited in
Males of H. higoensis can be distinguished from those of H. kikuyai by one of the embolus peaks being longer than the other (Fig.
Males (N = 11). Carapace yellow brown; opisthosoma light brown, with dark brown tergites close to each other; cheliceral groove with 11–13 denticles; 7 or 8 spinnerets. Measurements: BL 8.80–11.00, CL 4.40–5.25, CW 4.10–4.90, OL 4.40–5.60, OW 2.90–3.40; ALE > PLE > PME > AME; leg I 11.75 (3.45 + 1.50 + 2.60 + 2.70 + 1.50), leg II 12.40 (3.40 + 1.60 + 2.50 + 3.10 + 1.80), leg III 13.30 (3.20 + 1.60 + 2.40 + 3.80 + 2.30), leg IV 16.80 (4.40 + 1.20 + 3.30 + 5.20 + 2.70).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 43). Carapace and opisthosoma colour as in male; cheliceral groove with 11–16 pronounced denticles; tergites similar to male; 7–8 spinnerets. Measurements: BL 8.00–12.80, CL 4.30–6.10, CW 3.80–5.57, OL 4.10–6.50, OW 2.70–4.90; ALE > PLE > PME > AME; palp 8.20 (2.70 + 1.50 + 1.70 + 2.30), leg I 8.80 (2.80 + 1.75 + 1.70 + 1.55 + 1.00), leg II 8.93 (2.75 + 1.60 + 1.55 + 1.83 + 1.20), leg III 9.30 (2.70 + 1.60 + 1.50 + 2.20 + 1.30), leg IV 13.25 (3.65 + 1.80 + 2.40 + 3.40 + 2.00).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium (Fig.
We examined the male holotype of H. higoensis (Fig.
JAPAN · 1 ♂, 8 ♀♀; Kyushu, Kumamoto-ken, Hitoyoshi-shi, Fumotomachi, Hitoyoshi Ruins Park; 32.21N, 130.77E; alt. 140 m; 18 September 2013; D. Li and B. Wu leg.; XUX-2013-365 (♂ matured 19 July 2014 at CBEE), XUX-2013-360 to 364, 366 to 368 · 2 ♂♂, 5 ♀♀; Kyushu, Kumamoto-ken, Kumamoto-shi, Tatsutayama, Tatsuta National Park; 32.82N, 130.73E; alt. 60 m; 19 September 2013; D. Li and B. Wu leg.; XUX-2013-370 to 379 · 4 ♂♂, 4 ♀♀; Kyushu, Kumamoto-ken, Kumamoto-shi, Higashi-ku, Kozono 1-chome; 32.84N, 130.78E; alt. 100 m; 19 September 2013; D. Li and B. Wu leg.; XUX-2013-380 to 389 (XUX-2013-381, ♂ matured 2 August 2014 at CBEE) · 1 ♂, 3 ♀♀; Kyushu, Kumamoto-ken, Kumamoto-shi, Kasuga, Hanaokayama; 32.80N, 130.68E; alt. 120 m, 19 September 2013; D. Li and B. Wu leg.; XUX-2013-390 to 393 · 3 ♀♀; Miyazaki-ken, Nishiusuki-gun, Takachiho-cho, Mukoyama; 32.70N, 131.30E; alt. 320 m; 22 September 2013; D. Li and B. Wu leg.; XUX-2013-435 to 441 · 1 ♂, 3 ♀♀; Miyazaki-ken, Higashimorokata-gun, Kunitomi-cho, Honjo 11960-1; 32.00N, 131.34E; alt. 30 m; 23 September 2013; D. Li and B. Wu leg.; XUX-2013-456 (♂, matured 19 July 2014 at CBEE), XUX-2013-449 to 451 · 1 ♂, 12 ♀♀; Miyazaki-ken, Higashimorokata-gun, Kunitomi-cho, Honjo 4191; 31.98N, 131.33E; alt. 30 m; 23 September 2013; D. Li and B. Wu leg.; XUX-2013-457 to 467C · 1 ♀; Miyazaki-ken, Nishimorokata-gun, Takaharu-cho, Kamamuto, Lake Miike; 31.89N, 130.96E; alt. 360 m; 23 September 2013; D. Li and B. Wu leg.; XUX-2013-468 · 1 ♂, 4 ♀♀; 2 Kagoshima-ken, Kirishima-shi, Kirishima, Takachihokawara; 31.89N, 130.89E; alt. 960 m; 23 September 2013; D. Li and B. Wu leg.; XUX-2013-471 (♂, matured 8 June 2014 at CBEE), XUX-2013-472 to 474, 476.
The species is known from the following prefectures on the Japanese island Kyushu: Kumamoto-ken (Hitoyoshi-shi and Kumamoto-shi), Miyazaki-ken (Nishiusuki-gun, Higashimorokata-gun and Nishimorokata-gun), Kagoshima-ken (Kirishima-shi) (Fig.
Heptathela kikuyai
Ono, 1998: 16 (holotype: male, from Mt. Gozaga-dake, 20 km south of Oita-shi, Oita-ken, Kyushu, Japan, collected by N. Kikuya on 13 September 1979, deposited in
Males of H. kikuyai differ from those of H. higoensis by the two embolus peaks of a similar height, and the hooked tegular marginal apophysis (Fig.
Male and female genital anatomy of Heptathela kikuyai Ono, 1998 A, C 3409 (short for XUX-2013-409) B, D 3401 E–K 3405 A, B vulva dorsal view C, D vulva ventral view E palp prolateral view F palp ventral view G palp retrolateral view H–K palp distal view; 3405, 3409: Mt. Gozaga-dake, Oita-ken; 3401: Onomachi, Oita-ken. Scale bar: 0.5 mm.
Males (N = 5). Carapace and opisthosoma brown, with dark-spotted tergites; cheliceral groove with 10–12 denticles; 6 or 7 spinnerets. Measurements: BL 7.90–9.40, CL 4.00–5.05, CW 3.55–4.30, OL 3.90–4.60, OW 2.60–3.20; ALE > PLE > PME > AME; leg I 13.65 (3.90 + 1.90 + 2.75 + 3.30 + 1.80), leg II 13.35 (3.45 + 1.60 + 2.65 + 3.55 + 2.10), leg III 14.40 (3.30 + 1.50 + 2.75 + 4.45 + 2.40), leg IV 19.25 (4.85 + 1.90 + 3.35 + 6.00 + 3.15).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 30). Carapace and opisthosoma as in male; cheliceral groove with 12–17 pronounced denticles; tergites similar to male; 6–8 spinnerets. Measurements: BL 8.55–15.10, CL 4.30–7.90, CW 3.80–6.70, OL 4.40–8.75, OW 2.80–6.10; ALE > PLE > PME > AME; palp 9.15 (3.25 + 1.60 + 1.90 + 2.40), leg I 10.30 (3.30 + 1.90 + 1.90 + 2.00 + 1.20), leg II 10.15 (3.10 + 1.80 + 1.85 + 2.10 + 1.30), leg III 10.80 (3.00 + 1.80 + 1.80 + 2.60 + 1.60), leg IV 15.80 (4.50 + 2.20 + 2.80 + 4.10 + 2.20).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium. Paired receptacular clusters along the anterior margin of bursa copulatrix, indistinctly divided into two parts, with many small granules, inner ones with longer and slender genital stalks (Fig.
JAPAN · 6 ♀♀; Oita-ken, Bungoono-shi, Totoki, Onomachi; 33.06N, 131.52E; alt. 240 m; 20 September 2013; D. Li and B. Wu leg.; XUX-2013-394 to 396, 399 to 401 · 2 ♂♂, 4 ♀♀; Oita-ken, Bungoono-shi, Onomachi-Ando, Mt. Gozaga-dake; 33.11N, 131.55E; alt. 630 m; 20 September 2013; D. Li and B. Wu leg.; XUX-2013-404, 405, 408, 409, 411, 412 (405, ♂ matured 2 May 2014 at CBEE) · 2 ♂♂, 6 ♀♀; Oita-ken, Usuki-shi, Takeyama; 33.10N, 131.72E; alt. 80 m; 20 September 2013; D. Li and B. Wu leg.; XUX-2013-414, 416 to 422 · 1 ♂, 8 ♀♀; Oita-ken, Usuki-shi, Inukai-machi Sasamuta; 33.09N, 131.67E; alt. 120 m; 21 September 2013; D. Li and B. Wu leg.; XUX-2013-424 to 430, 432, 433 · 6 ♀♀; Miyazaki-ken, Nishiusuki-gun, Takachiho-cho, Iwato; 32.73N, 131.35E; alt. 400 m; 22 September 2013; D. Li and B. Wu leg.; XUX-2013-442 to 444, 446 to 448.
The species is known from the following prefectures on the Japanese island Kyushu: Oita-ken (Bungoono-shi and Usuki-shi) and Miyazaki-ken (Nishiusuki-gun) (Fig.
Liphistius kimurai
Kishida, 1920: 362 (holotype: female, from Shiroyama, Kagoshima, Kyushu, Japan, collected by A. Kimura in October 1920, lost in the Science College Museum of the Tokyo Imperial University (Haupt, 1983); neotype: male, from the same locality as for the original type specimen, collected by J. Haupt on 21 March 1982, matured in August 1982, deposited in ZMH, but the neotype may be lost according to
Heptathela kimurai
Kishida, 1923: 236;
Females of H. kimurai resemble H. higoensis females but differ by a slightly curved dorso-posterior margin of the genital area (Fig.
Females (N = 9). Carapace yellow brown; opisthosoma brown, with brown tergites close to each other; cheliceral groove with 12–14 vestigial denticles; seven spinnerets. Measurements: BL 9.30–13.80, CL 4.88–6.30, CW 4.10–5.50, OL 4.80–7.30, OW 3.45–5.40; ALE > PLE > PME > AME; palp 7.77 (2.57 + 1.50+ 1.60 + 2.10), leg I 9.15 (2.85 + 1.70 + 1.75 + 1.65 + 1.20), leg II 8.88 (2.65 + 1.65 + 1.55 + 1.88 + 1.15), leg III 9.63 (2.70 + 1.70 + 1.60 + 2.30 + 1.33), leg IV 14.08 (3.88 + 1.90 + 2.50 + 3.60 + 2.20).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium. Paired of receptacular clusters along the anterior margin of bursa copulatrix, divided into two parts, the inner main part forming a large granulate tubercle, the outer part with several small granules (Fig.
Male: unknown.
We could not examine the presumably lost neotype male (
JAPAN · 9 ♀♀; Kagoshima-ken, Kagoshima-shi, Shiroyama-cho, Shiroyama Park; 31.60N, 130.55E; alt. 100 m; 18 September 2013; D. Li and B. Wu leg.; XUX-2013-349, 351, 352, 354 to 359.
The species is known from the Kagoshima prefecture on the Japanese island Kyushu (Fig.
Heptathela yakushimaensis
Ono, 1998: 23 (holotype: female, from Mt. Kunibaidake, Yakushima Island, Kagoshima-ken, Japan, collected by A. Tanikawa on 15 July 1990, deposited in
Heptathela kimurai yakushimaensis:
Males of H. yakushimaensis differ from those of all other Kyushu group Heptathela species by a strongly serrated prolateral conductor margin (Fig.
Males (N = 2). Carapace brown; opisthosoma light brown, with dark brown tergites close to each other; cheliceral groove with 10–13 denticles; 7 spinnerets. Measurements: BL 8.70–10.50, CL 4.27–5.50, CW 4.00–4.90, OL 3.80–6.10, OW 2.40–4.40; ALE > PLE > PME > AME; leg I 13.27 (3.70 + 1.60 + 2.72 + 3.50 + 1.75), leg II 13.71 (3.55 + 1.65 + 2.80 + 3.75 + 1.96), leg III 14.65 (3.52 + 1.65 + 2.78 + 4.50 + 2.20), leg IV 18.64 (4.35 + 1.70 + 3.72 + 6.20 + 2.67).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 8). Carapace and opisthosoma colour as in male; cheliceral groove with 12–15 pronounced denticles; tergites similar to male; 7 spinnerets. Measurements: BL 9.10–14.80, CL 4.50–6.80, CW 4.05–5.90, OL 4.70–8.60, OW 3.00–6.40; ALE > PLE > PME > AME; palp 9.72 (3.22 + 1.70 + 2.10 + 2.70), leg I 11.15 (3.45 + 1.90 + 2.05 + 2.35 + 1.40), leg II 9.61 (3.07 + 1.90 + 1.00 + 2.36 + 1.28), leg III 11.75 (3.25 + 1.95 + 2.05 + 2.80 + 1.70), leg IV 16.30 (4.25 + 2.25 + 2.90 + 4.40 + 2.50).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium. A pair of receptacular clusters along the anterior margin of bursa copulatrix, divided into two parts, the inner part is similar or smaller than the outer part, on which there are several small granules (Fig.
JAPAN · 2 ♂♂, 8 ♀♀; Kagoshima-ken, Kumage-gun, Yakushima-cho, Mt. Kankake-dake; 30.37N, 130.39E; alt. 170 m; 24 September 2013; D. Li and B. Wu leg.; XUX-2013-490, 491, 493 to 500 (500, ♂ matured 2 August 2014 at CBEE).
The species is known from the Kagoshima prefecture on the Japanese island Yakushima (Fig.
Diagnosis. The males of the Amami group differ from those of the other two groups by the rugose conductor with a spiniform apex, the contrategulum with a strongly serrated margin whereas it is nearly rectangular in Kyushu group and weakly serrated in Okinawa group, and the embolus with a wide and flat opening (Fig.
Male and female genital anatomy of Heptathela amamiensis Haupt, 1983 A, C 3278 (short for XUX-2013-278) B, D, E 3285 F–L 3283 A, B vulva dorsal view C, D vulva ventral view E vulva distal view F right palp prolateral view G right palp ventral view H right palp retrolateral view I–L left palp distal view. Scale bar: 0.5 mm.
Monophyly. The Bayesian analyses based on concatenated two genes and two partitions (for details, see
Composition. H. amamiensis Haupt, 1983, H. kanenoi Ono, 1996, H. kojima sp. nov., H. sumiyo sp. nov., and H. uken sp. nov.
Distribution. Amamioshima and Tokunoshima (Fig.
Heptathela kimurai amamiensis
Haupt, 1983: 283 (holotype: female, from Naze, Amami-oshima, Japan, collected by J. Haupt on 26 March 1980, deposited in ZMH, holotype presumably lost (
Males of H. amamiensis differ from those of H. sumiyo sp. nov. by the wider saddle-shaped embolus in the prolateral view, and the narrower conductor base in the ventral view (Fig.
Male. Carapace brown; opisthosoma light brown, with dark brown tergites; cheliceral groove with 13 denticles; 8 spinnerets. Measurements: BL 12.85, CL 6.50, CW 5.98, OL 6.65, OW 4.00; ALE > PLE > PME > AME; leg I 18.25 (5.00 + 2.45 + 3.80 + 4.60 + 2.40), leg II 18.75 (4.85 + 2.40 + 3.75 + 5.00 + 2.75), leg III 20.10 (4.90 + 2.60 + 3.60 + 5.90 + 3.10), leg IV 25.30 (6.00 + 2.60 + 5.10 + 7.70 + 3.90).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 5). Carapace and opisthosoma colour as in male; cheliceral groove with 12–14 pronounced denticles; tergites similar to those of male; seven or eight spinnerets. Measurements: BL 11.08–16.90, CL 5.45–6.90, CW 4.70–6.20, OL 6.61–9.10, OW 4.70–7.38; ALE > PLE > PME > AME; palp 12.36 (4.12 + 2.11 + 2.75 + 3.38), leg I 14.12 (4.52 + 2.50 + 2.70 + 2.83 + 1.57), leg II 14.06 (4.25 + 2.37 + 2.61 + 3.15 + 1.68), leg III 15.13 (4.27 + 2.56 + 2.55 + 3.55 + 2.20), leg IV 21.86 (5.84 + 2.91 + 4.08 + 5.81 + 3.22).
Female genitalia. A pair of indistinct depressions on the ventro-lateral part of genital atrium (Fig.
JAPAN · 2 ♂♂, 5 ♀♀; Kagoshima-ken, Amami-Oshima, Amami-shi, Nazehirata-cho, Michinoshima Loop Bridge; 28.36N, 129.50E; alt. 60 m; 15 September 2013; D. Li and B. Wu leg.; XUX-2013-276, 278, 281 to 285.
The species is known from the Japanese island Amamioshima (Fig.
Heptathela kanenoi
Ono, 1996: 158 (holotype: male, from Mikyo, Amagi-cho, Tokunoshima, Kagoshima-ken, Japan, collected by M. Owada and S. Kaneno on 2 November 1992, deposited in
Males of H. kanenoi can be distinguished from those of all other Amami group Heptathela species by lacking a spiniform conductor apex (Fig.
Males (N = 6). Carapace yellow brown; opisthosoma brown, with dark-spotted tergites close to each other; cheliceral groove with 10–13 vestigial denticles; 7 spinnerets. Measurements: BL 9.80–11.60, CL 5.10–6.00, CW 4.60–5.40, OL 5.15–5.60, OW 4.00–4.85; ALE > PLE > PME > AME; leg I 17.85 (4.80 + 2.30 + 3.80 + 4.60 + 2.35), leg II 18.90 (4.90 + 2.30 + 3.80 + 5.20 + 2.70), leg III 20.80 (4.70 + 2.30 + 4.00 + 6.50 + 3.30), leg IV 26.00 (6.20 + 2.50 + 5.10 + 8.20 + 4.00).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 11). Carapace and opisthosoma colour as in male; cheliceral groove with 12–14 pronounced denticles; tergites similar to male; seven or eight spinnerets. Measurements: BL 8.30–12.90, CL 4.30–6.28, CW 3.60–5.40, OL 4.38–6.50, OW 3.70–6.20; ALE > PLE > PME > AME; palp 7.00 (2.50 + 1.35 + 1.55 + 1.60), leg I 7.75 (2.30 + 1.25 + 1.60 + 1.50 + 1.10), leg II 7.60 (2.60 + 1.10 + 1.50 + 1.50 + 0.90), leg III 8.35 (2.45 + 1.50 + 1.10 + 2.00 + 1.30), leg IV 12.15 (3.40 + 1.50 + 2.25 + 3.20 + 1.80).
Female genitalia. A pair of indistinct depressions on the ventro-lateral part of genital atrium. Paired receptacular clusters along the anterior margin of bursa copulatrix, divided into two parts, the inners larger than the outers, without genital stalks (Fig.
JAPAN · 1 ♂, 1 ♀; Kagoshima-ken, Tokunoshima, Amagi-cho, Mikyo; 27.77N, 128.95E; alt. 180 m; 16 September 2013; D. Li and B. Wu leg.; XUX-2013-315 to 316 · 3 ♂♂, 7 ♀♀; Kagoshima-ken, Oshima-gun, Tokunoshima, Tokunoshima-cho, Tokuwase; 27.79N, 129.01E; alt. 150 m; 17 September 2013; D. Li and B. Wu leg.; XUX-2013-323 to 332 · 2 ♂♂, 3 ♀♀; Kagoshima-ken, Tokunoshima, Amagi-cho, Mikyo; 27.77N, 128.95E; alt. 130 m; 17 September 2013; D. Li and B. Wu leg.; XUX-2013-333 to 338.
The species is endemic to the Japanese island Tokunoshima (Fig.
Holotype : JAPAN · ♂; Kagoshima-ken, Oshima-gun, Tokunoshima, Isen-cho, Kojima; 27.74N, 128.91E; alt. 160 m; 17 September 2013; D. Li and B. Wu leg.; XUX-2013-346 (matured 10 October 2013 at CBEE).
Paratypes : JAPAN · 2 ♂♂, 6 ♀♀; same data as for holotype; XUX-2013-339, 340, 342 to 345, 347, 348.
Males of H. kojima sp. nov. differ from those of H. amamiensis and H. kanenoi by a wide leaf-shaped conductor (Fig.
Male and female genital anatomy of Heptathela kojima sp. nov. A, C 3339 (short for XUX-2013-339) B, D 3340 E–G 3346 (holotype) H–K 3344 A, B vulva dorsal view C, D vulva ventral view E palp prolateral view F palp ventral view G palp retrolateral view H–K palp distal view. Scale bar: 0.5 mm.
Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites close to each other; cheliceral groove with eleven denticles; seven spinnerets. Measurements: BL 7.30, CL 3.60, CW 3.30, OL 3.60, OW 2.70; ALE > PLE > PME > AME; leg I 14.90 (4.20 + 165 + 3.15 + 4.00 + 1.90), leg II 16.10 (4.20 + 1.85 + 3.25 + 4.40 + 2.40), leg III 16.85 (4.25 + 1.90 + 3.20 + 5.00 + 2.50), leg IV 21.70 (5.50 + 2.00 + 4.20 + 6.90 + 3.10).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 6). Carapace and opisthosoma colour as in male; cheliceral groove with 13 pronounced denticles; tergites similar to male; 6–8 spinnerets. Measurements: BL 9.00–11.50, CL 4.90–6.15, CW 4.25–5.20, OL 4.30–6.10, OW 3.00–4.50; ALE > PLE > PME > AME; palp 9.05 (3.25 + 1.65 + 1.85 + 2.30), leg I 10.50 (3.35 + 1.80 + 2.00 + 2.10 + 1.25), leg II 10.05 (3.10 + 1.80 + 1.75 + 2.10 + 1.30), leg III 10.45 (2.90 + 1.95 + 1.65 + 2.40 + 1.55), leg IV 15.20 (4.40 + 2.20 + 2.60 + 4.00 + 2.00).
Female genitalia. A pair of indistinct depressions on the ventro-lateral part of genital atrium (Fig.
The species epithet, a noun in apposition, refers to the type locality.
The species is endemic to the Japanese island Tokunoshima (Fig.
Holotype : JAPAN · ♂; Kagoshima-ken, Amami-Oshima, Amami-shi, Sumiyo-cho, Santaro-toge Pass; 28.28N, 129.42E; alt. 360 m; 15 September 2013; D. Li and B. Wu leg.; XUX-2013-293.
Paratypes : JAPAN · 4 ♂♂, 6 ♀♀; same data as for holotype; XUX-2013-287 to 292, 294 to 296B.
Males of H. sumiyo sp. nov. can be distinguished from those of H. kanenoi by the spiniform conductor apex (Fig.
Male and female genital anatomy of Heptathela sumiyo sp. nov. A–C 3293 (holotype, short for XUX-2013-293) D–G 3292 H, J 3288 I, K 3296 A palp prolateral view B palp ventral view C palp retrolateral view D–G palp distal view H, I vulva dorsal view J, K vulva ventral view. Scale bar: 0.5 mm.
Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; cheliceral groove with eleven denticles; seven spinnerets. Measurements: BL 15.40, CL 7.70, CW 6.80, OL 8.00, OW 5.55; ALE > PLE > PME > AME; leg I 23.05 (6.05 + 3.10 + 4.90 + 6.10 + 2.90), leg II 24.46 (6.48 + 3.08 + 5.05 + 6.80 + 3.05), leg III 26.36 (6.10 + 3.20 + 5.05 + 8.20 + 3.81), leg IV 32.70 (8.00 + 3.50 + 6.75 + 9.70 + 4.75).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 6). Carapace and opisthosoma colour as in male; cheliceral groove with 13 or 14 pronounced denticles; tergites similar to those of male; seven or eight spinnerets. Measurements: BL 11.70–15.60, CL 6.25–7.55, CW 5.40–6.43, OL 5.40–8.80, OW 4.00–7.35; ALE > PLE > PME > AME; palp 12.90 (4.50 + 2.25 + 2.70 + 3.45), leg I 14.44 (4.55 + 2.68 + 2.55 + 3.08 + 1.58), leg II 13.93 (4.09 + 2.49 + 2.35 + 3.20 + 1.80), leg III 15.61 (4.40 + 2.70 + 2.68 + 3.80 + 2.03), leg IV 22.45 (6.20 + 3.20 + 4.15 + 6.10 + 2.80).
Female genitalia. A pair of indistinct depressions on the ventro-lateral part of genital atrium (Fig.
The species epithet, a noun in apposition, refers to the type locality.
The species is known from the Japanese island Amamioshima (Fig.
Holotype : JAPAN · ♂; Kagoshima-ken, Amami, Uken-son, Oshima-gun, Road No. 85, Redsoil Park; 28.24N, 129.34E; alt. 260 m; 15 September 2013; D. Li and B. Wu leg.; XUX-2013-297.
Paratypes : JAPAN · 2 ♂♂, 2 ♀♀; same data as for holotype; XUX-2013-298, 301, 302, 304 · 3 ♂♂, 5 ♀♀; Kagoshima-ken, Amami, Yamato-son, Amami Forest Park; 28.31N, 129.33E; alt. 300 m; 17 September 2013; D. Li and B. Wu leg.; XUX-2013-305 to 314.
Males of H. uken sp. nov. can be distinguished from those of H. kanenoi by the spiniform conductor apex (Fig.
Male and female genital anatomy of Heptathela uken sp. nov. A–G 3297 (holotype, short for XUX-2013-297) H, K, J 3301 I, L, M 3309 A palp prolateral view B palp ventral view C palp retrolateral view D–G palp distal view H, I vulva dorsal view K, L vulva ventral view J, M vulva distal view; 3297, 3301: Uken-son, Amamioshima; 3309: Yamato-son, Amamioshima. Scale bar: 0.5 mm.
Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; cheliceral groove with eight denticles; seven spinnerets. Measurements: BL 11.60, CL 6.10, CW 5.60, OL 6.00, OW 4.30; ALE > PLE > PME > AME; leg I 17.47 (4.80 + 2.28 + 3.67+ 4.42 + 2.30), leg II 17.95 (4.65 + 2.30 + 3.60 + 4.90 + 2.50), leg III 18.80 (4.45 + 2.40 + 3.55 + 5.20 + 3.20), leg IV 23.60 (5.60 + 2.45 + 4.45 + 7.20 + 3.90).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 8). Carapace and opisthosoma colour as in male; cheliceral groove with 13–15 pronounced denticles; tergites similar to male; 7 or 8 spinnerets. Measurements: BL 11.80–16.00, CL 5.90–8.26, CW 5.20–7.20, OL 6.00–8.10, OW 4.60–6.00; ALE > PLE > PME > AME; palp 10.04 (3.05 + 1.89 + 2.30 + 2.80), leg I 12.05 (3.90 + 2.10 + 2.25 + 2.40 + 1.40), leg II 11.30 (3.55 + 2.05 + 1.80+ 2.45 + 1.45), leg III 12.42 (3.60 + 2.05 + 2.10 + 3.00 + 1.67), leg IV 18.33 (5.15 + 2.40 + 3.30 + 4.78 + 2.70).
Female genitalia. A pair of indistinct depressions on the ventro-lateral part of genital atrium (Fig.
The species epithet, a noun in apposition, refers to the type locality.
The species is known from the Japanese island Amamioshima (Fig.
Diagnosis. The males of the Okinawa group differ from those of the Kyushu group by the semi-elliptic contrategulum (Fig.
Male and female genital anatomy of Heptathela yanbaruensis Haupt, 1983 A, C 2315 (short for XUX-2012-315) B, D 2441 E–G 2322B H–K 2311 A, B vulva dorsal view C, D vulva ventral view E palp prolateral view F palp ventral view G palp retrolateral view H–K palp distal view; 2311, 2315, 2322B: Yona, Okinawajima; 2441: Taiho Dam, Okinawajima. Scale bar: 0.5 mm.
Monophyly. The Bayesian analyses based on concatenated two genes and two partitions (for details, see
Composition. H. yanbaruensis Haupt, 1983, H. aha sp. nov., H. gayozan sp. nov., H. kubayama sp. nov., H. mae sp. nov., H. otoha sp. nov., H. shuri sp. nov., H. tokashiki sp. nov., H. unten sp. nov., and H. crypta sp. nov.
Distribution. Okinawajima, Iheyajima, Tokashikijima (Fig.
Heptathela kimurai yanbaruensis
Haupt, 1983: 284 (holotype: male, from Yona, Okinawa, Japan, collected by J. Haupt on 15 April 1977, deposited in ZMH, where the type may be lost (
Males of H. yanbaruensis can be distinguished from those of H. helios by the contrategulum that is distinctly curved in the middle (Fig.
Males (N = 3). Carapace yellow brown; opisthosoma light brown, with dark-spotted tergites close to each other; cheliceral groove with 10–12 denticles; seven spinnerets. Measurements: BL 8.53–9.22, CL 4.21–4.57, CW 3.71–3.75, OL 4.20–4.48, OW 2.30–2.80; ALE > PLE > PME > AME; leg I 14.28 (4.15 + 1.70 + 3.05 + 3.56 + 1.82), leg II 15.15 (3.97 + 1.70 + 3.07 + 4.13 + 2.28), leg III 15.24 (4.00 + 1.68 + 3.08 + 4.10 + 2.38), leg IV 19.78 (4.75 + 1.37 + 3.91 + 6.45 + 3.30).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 15). Carapace and opisthosoma colour as in male; cheliceral groove with 11–14 pronounced denticles; tergites similar to male; seven spinnerets. Measurements: BL 7.78–10.33, CL 3.90–5.10, CW 3.40–4.23, OL 3.90–6.00, OW 2.80–4.70; ALE > PLE > PME > AME; palp 8.70 (2.96 + 1.51 + 2.11 + 2.12), leg I 10.20 (3.12 + 1.75 + 1.93 + 2.12 + 1.28), leg II 9.97 (3.05 + 1.72 + 1.77 + 2.13 + 1.30), leg III 10.05 (2.91 + 1.78 + 1.48 + 2.47 + 1.41), leg IV 14.77 (4.08 + 2.10 + 2.51 + 3.97 + 2.11).
Female genitalia. A pair of depressions on the ventro-lateral part of the genital atrium (Fig.
JAPAN · 3 ♂♂, 9 ♀♀; Okinawa-ken, Kunigami-son, Yona, Tropical Biosphere Research Centre field station, University of the Ryukyu; 26.76N, 128.22E; alt. 20 m; 18 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-310 to 322B · 6 ♀♀; Okinawa-ken, Ogimi-son, Taiho Dam; 26.65N, 128.16E; alt. 80 m; 24 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-441 to 446.
The species is endemic to the Japanese island Okinawajima (Fig.
Holotype : JAPAN · ♀; Okinawa-ken, Iheyajima Island, Mt. Aha-dake; 27.02N, 127.93E; alt. 10 m; 26 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-502.
Paratypes : JAPAN · 6 ♀♀; same data as for holotype; XUX-2012-504 to 519.
Females of H. aha sp. nov. cannot be distinguished morphologically from those of H. gayozan sp. nov. but can be distinguished from those of H. kubayama sp. nov. by the receptacular clusters without genital stalks; and from those of H. mae sp. nov. by the inner receptacular clusters similar to or larger than laterals (Fig.
Female (Holotype). Carapace brown; opisthosoma brown, brown tergites with black plaques; cheliceral groove with 12 pronounced denticles; seven spinnerets. Measurements: BL 12.60, CL 5.55, CW 5.00, OL 6.82, OW 5.50; ALE > PLE > PME > AME; palp 9.90 (3.40 + 1.71 + 2.18 + 2.61), leg I 11.34 (3.31 + 2.05 + 2.31 + 2.35 + 1.32), leg II 11.65 (3.51 + 1.98 + 2.03 + 2.53 + 1.60), leg III 11.92 (3.30 + 1.90 + 2.05 + 2.97 + 1.70), leg IV 16.96 (4.88 + 2.22 + 2.98 + 34.45 + 2.43).
Female genitalia. A pair of depressions on the ventro-lateral part of the genital atrium (Fig.
Male. Unknown.
The species epithet, a noun in apposition, refers to the type locality.
The species is endemic to the Japanese island Iheyajima (Fig.
Holotype : JAPAN · ♀; Okinawa-ken, Iheyajima Island, Mt. Gayozan; 27.02N, 127.97E; alt. 25 m; 27 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-511.
Paratypes : JAPAN · 2 ♀♀; same data as for holotype; XUX-2012-513, 515.
Females H. gayozan sp. nov. cannot be distinguished morphologically from those of H. aha sp. nov. but can be distinguished from those of H. kubayama sp. nov. by the receptacular clusters without genital stalks; and from those of H. mae sp. nov. by the inner receptacular clusters that are equal in size, or slightly larger than laterals (Fig.
Female (holotype). Carapace brown; opisthosoma light brown, with dark brown tergites close to each other; cheliceral groove with 12 denticles; seven spinnerets. Measurements: BL 10.00, CL 4.40, CW 3.88, OL 4.84, OW 3.48; ALE > PLE > PME > AME; palp 7.32 (2.72 + 1.50 + 1.80 + 1.30), leg I 9.61 (2.95 + 1.62 + 1.82 + 1.97 + 1.25), leg II 9.38 (2.80 + 1.58 + 1.71 + 2.07 + 1.22), leg III 9.60 (2.65 + 1.55 + 1.62 + 2.28 + 1.50), leg IV 13.89 (3.75 + 1.82 + 2.52 + 3.75 + 2.05).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium (Fig.
Male. Unknown.
The species epithet, a noun in apposition, refers to the type locality.
The species is endemic to the Japanese island Iheyajima (Fig.
Holotype : JAPAN · ♀; Okinawa-ken, Iheyajima Island, Mt. Kubayama Nature Conservation Area; 27.09N, 128.02E; alt. 85 m; 26 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-486.
Paratypes : JAPAN · 8 ♀♀; same data as for holotype; XUX-2012-479, 481 to 485, 487 to 488.
Females of H. kubayama sp. nov. can be distinguished from those of H. gayozan sp. nov. and H. mae sp. nov. by paired receptacular clusters with short genital stalks (Fig.
Female (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites close to each other; cheliceral groove with 12 pronounced denticles; six spinnerets. Measurements: BL 10.20, CL 4.89, CW 4.18, OL 4.90, OW 3.60; ALE > PLE > PME > AME; palp 8.25 (2.82 + 1.45 + 1.80 + 2.18), leg I 9.66 (3.10 + 1.70 + 1.81 + 1.90 + 1.15), leg II 8.90 (2.90 + 1.55 + 1.75 + 1.70 + 1.00), leg III 9.59 (2.81 + 1.65 + 1.60 + 2.13 + 1.40), leg IV 13.67 (4.25 + 2.00 + 2.00 + 3.54 + 1.88).
Female genitalia. A pair of depressions on the ventro-lateral part of the genital atrium (Fig.
Male. Unknown.
The species epithet, a noun in apposition, refers to the type locality.
The species is endemic to the Japanese island Iheyajima (Fig.
Holotype : JAPAN · ♀; Okinawa-ken, Iheyajima Island, Mt. Mae-dake; 27.06N, 127.99E; alt. 10 m; 26 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-497.
Paratypes : JAPAN · 5 ♀♀; same data as for holotype; XUX-2012-494 to 496, 498, 500 · 3♀♀; Okinawa Prefecture, Iheyajima Island, Mt. Mae-dake; 27.06N, 127.99E; alt. 20 m; 10 May 2014; D. Li and B. Wu leg.; XUX-2014-079A to 079C.
Females of H. mae sp. nov. can be distinguished from those of H. aha sp. nov. and H. gayozan sp. nov. by the lateral receptacular clusters being larger than the inner ones (Fig.
Female (Holotype). Carapace brown; opisthosoma brown, brown tergites with black plaques; cheliceral groove with 12 denticles; seven spinnerets. Measurements: BL 9.28, CL 4.48, CW 4.00, OL 5.12, OW 3.78; ALE > PLE > PME > AME; palp 7.80 (2.70 + 1.32 + 1.71 + 2.07), leg I 9.25 (3.00 + 1.52 + 1.80 + 1.78 + 1.15), leg II 8.55 (2.68 + 1.20 + 1.72 + 1.75 + 1.20), leg III 9.61 (2.75 + 1.60 + 1.55 + 2.31 + 1.40), leg IV 13.41 (3.90 + 1.70 + 2.31 + 3.60 + 1.90).
Female genitalia. A pair of depressions on the ventro-lateral part of the genital atrium (Fig.
Male. Unknown.
The species epithet, a noun in apposition, refers to the type locality.
The species is endemic to the Japanese island Iheyajima (Fig.
Holotype : JAPAN · ♀; Okinawa-ken, Nakijin-son, Mt. Otoha-dake; 26.67N, 127.97E; alt. 80 m; 27 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-535.
Paratypes : JAPAN · 3 ♀♀; Okinawa Prefecture, Motobu-cho, Yamazato; 26.67N, 127.91E; alt. 160 m; 11 May 2014; D. Li and B. Wu leg.; XUX-2014-094, 097, 099 · 3 ♀♀; Okinawa Prefecture, Nakijin-son, Mt. Otoha-dake; 26.67N, 127.97E; alt. 100 m; 11 May 2014; D. Li and B. Wu leg.; XUX-2014-101, 102, 103.
Females of H. otoha sp. nov. can be distinguished from those of H. yanbaruensis, H. unten sp. nov., and H. crypta sp. nov. by the inner receptacular clusters with several granules, albeit these inner clusters are difficult to be separated from the laterals (Fig.
Female (Holotype). Carapace yellow brown; opisthosoma light brown, with brown and black-spotted tergites; cheliceral groove with 13 pronounced denticles; seven spinnerets. Measurements: BL 10.92, CL 4.75, CW 4.35, OL 6.60, OW 4.90; ALE > PLE > PME > AME; palp 8.30 (2.78 + 1.39 + 1.90 + 2.23), leg I 9.80 (3.05 + 1.65 + 1.95 + 1.95 + 1.20), leg II 9.21 (2.90 + 1.50 + 1.81 + 1.90 + 1.10), leg III 9.84 (2.70 + 1.70 + 1.62 + 2.35 + 1.47), leg IV 14.74 (4.00 + 1.90 + 2.70 + 4.03 + 2.11).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium (Fig.
Male. Unknow.
The species epithet, a noun in apposition, refers to the type locality.
The species is endemic to the Japanese island Okinawajima (Fig.
Holotype : JAPAN · ♀; Okinawa-ken, Naha, Shuri, Sueyoshi Park; 26.23N, 127.72E; alt. 45 m; 17 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-309.
Paratype : JAPAN · 1 ♀; same data as for holotype; XUX-2012-308.
Females of H. shuri sp. nov. can be distinguished from those of H. tokashiki sp. nov. by paired receptacular clusters with larger granules (Fig.
Female (Holotype). Carapace yellow brown; opisthosoma brown, tergites with brown plaques; cheliceral groove with 12 pronounced denticles; seven spinnerets. Measurements: BL 10.78, CL 5.30, CW 4.12, OL 6.25, OW 4.40; ALE > PLE > PME > AME; palp 8.93 (3.15 + 1.43 + 2.03 + 2.32), leg I 10.91 (3.48 + 1.87 + 2.05 + 2.21 + 1.30), leg II 10.64 (3.18 + 1.85 + 1.86 + 2.28 + 1.47), leg III 11.26 (3.03 + 1.81 + 1.77 + 2.90 + 1.75), leg IV 17.02 (4.62 + 2.28 + 3.07 + 4.50 + 2.55).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium (Fig.
Male. Unknown.
The species epithet, a noun in apposition, refers to the type locality.
The species is endemic to the Japanese island Okinawajima (Fig.
Holotype : JAPAN · ♀; Okinawa-ken, Tokashikijima Island, Aharen; 26.19N, 127.37E; alt. 100 m; 8 May 2014; D. Li and B. Wu leg.; XUX-2014-062.
Paratypes : JAPAN · 25 ♀♀; same data as for holotype; XUX-2014-046 to 063D · 3 ♀♀; same data as for holotype; 27 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-417, 421, 425.
Females of H. tokashiki sp. nov. can be distinguished from H. shuri sp. nov. by the long inner receptacular clusters with several granules (Fig.
Female (Holotype). Carapace brown; opisthosoma dark brown, with dark brown tergites close to each other; cheliceral groove with eleven pronounced denticles; seven spinnerets. Measurements: BL 11.80, CL 5.15, CW 4.10, OL 6.80, OW 5.10; ALE > PLE > PME > AME; palp 8.60 (3.05 + 1.50 + 1.95 + 2.10), leg I 10.50 (3.35 + 1.85 + 2.15 + 2.10 + 1.05), leg II 10.45 (3.40 + 1.75 + 2.00 + 2.20 + 1.10), leg III 10.95 (3.20 + 1.85 + 1.80 + 2.60 + 1.50), leg IV 15.70 (4.60 + 2.00 + 2.80 + 3.90 + 2.40).
Female genitalia. A pair of depressions on the ventro-lateral part of the genital atrium (Fig.
Male. Unknown.
The species epithet, a noun in apposition, refers to the type locality.
The species is endemic to the Japanese island Tokashikijima (Fig.
Holotype : JAPAN · ♂; Okinawa-ken, Nakijin-son, Unten Port; 26.68N, 128.00E; alt. 25 m; 27 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-522.
Paratypes : JAPAN · 1 ♂, 2 ♀♀; same data as for holotype; XUX-2012-523, 527, 528A · 3 ♀♀; same data as for holotype; 10 May 2014; D. Li and B. Wu leg.; XUX-2014-083, 083A, 083B.
Males of H. unten sp. nov. can be distinguished from those of H. yanbaruensis by the blunt tegular marginal apophysis (Fig.
Male genital anatomy of Heptathela unten sp. nov. and H. crypta sp. nov. A–D Heptathela unten sp. nov., 2522 (holotype, short for XUX-2012-522) E–K H. crypta sp. nov. E–H 2328 (holotype) I–K 2460 A, I prolateral view B, K retrolateral view J ventral view C–H distal view; 2522: Unten Port, Okinawajima; 2328: Taira, Haneiji–Dam, Okinawajima; 2460: Yofuke, Okinawajima. Scale bar: 0.5 mm.
Female genital anatomy of Heptathela unten sp. nov. and H. crypta sp. nov. A, D 2527 B, E 4083A C, F 4027B G, J 2324 H, K 2327 I, L 2457 A–C, G–I dorsal view D–F, J–L ventral view; 2527, 4083A: Unten Port, Okinawajima; 4027B: Mt. Nago-dake, Okinawajima; 2324, 2327: Taira, Haneiji-Dam, Okinawajima; 2457: Yofuke, Okinawajima. Scale bar: 0.5 mm.
Male (Holotype). Carapace yellow brown; opisthosoma brown, with dark brown tergites close to each other; cheliceral groove with ten denticles; seven spinnerets. Measurements: BL 10.00, CL 5.05, CW 4.60, OL 4.60, OW 2.90; ALE > PLE > PME > AME; leg I 15.71 (4.48 + 1.93 + 3.30 + 4.00 + 2.00), leg II 16.59 (4.45 + 1.90 + 3.37 + 4.62 + 2.25), leg III 18.29 (4.50 + 1.93 + 3.58 + 5.53 + 2.75), leg IV 23.40 (5.82 + 2.28 + 4.52 + 7.30 + 3.48).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 5). Carapace and opisthosoma colour as in male, dark brown tergites separated from each other; cheliceral groove with 12 or 13 pronounced denticles; seven or eight spinnerets. Measurements: BL 7.81–12.00, CL 3.60–4.55, CW 3.30–4.40, OL 4.10–7.60, OW 3.00–6.20; ALE > PLE > PME > AME; palp 6.64 (2.19 + 1.22 + 1.45 + 1.78), leg I 7.48 (2.47 + 1.35 + 1.22 + 1.52 + 0.92), leg II 7.40 (2.21 + 1.30 + 1.21 + 1.65 + 1.03), leg III 7.79 (2.11 + 1.38 + 1.20 + 1.90 + 1.20), leg IV 11.55 (3.20 + 1.55 + 2.10 + 3.05 + 1.65).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium indistinct (Fig.
The species epithet, a noun in apposition, refers to the type locality, Unten Port.
The species is endemic to the Japanese island Okinawajima (Fig.
Holotype : JAPAN · ♂; Okinawa-ken, Nago-shi, Haneiji-Dam, Taira; 26.59N, 128.03E; alt. 100 m; 18 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-328.
Paratypes : JAPAN · 1 ♂, 3 ♀♀; same data as for holotype; XUX-2012-324, 326, 327, 333A · 2 ♂♂, 4 ♀♀; Okinawa Prefecture, Nago-shi, County Road 18 south, Nago/Yofuke; 26.57N, 128.01E; alt. 150 m; 24 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-457 to 462 · 3 ♀♀; Okinawa Prefecture, Nago-shi, Mt. Nago-dake; 26.58N, 128.01E; alt. 220 m; 06 May 2014; D. Li and B. Wu leg.; XUX-2014-027 to 027B.
Males and females of H. crypta sp. nov. cannot be distinguished morphologically from H. unten sp. nov. (Figs
Male (Holotype). Carapace and opisthosoma description see H. unten sp. nov.; cheliceral groove with nine denticles of variable size; seven spinnerets. Measurements: BL 7.88, CL 4.01, CW 3.51, OL 4.23, OW 3.18; ALE > PLE > PME > AME; leg I 10.15 (3.60 + 1.50 + 2.38 + 1.00 + 1.67), leg II 13.08 (3.48 + 1.58 + 2.49 + 3.53 + 2.00), leg III 14.27 (3.38 + 1.55 + 2.65 + 4.30 + 2.39), leg IV 18.19 (4.45 + 1.63 + 3.50 + 5.50 + 3.11).
Palp. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 10). Carapace and opisthosoma description see H. unten sp. nov.; chelicerae with promargin of cheliceral groove with 13–14 pronounced denticles of variable size; seven spinnerets. Measurements: BL 8.35–16.50, CL 4.07–5.10, CW 3.30–4.80, OL 4.70–6.80, OW 3.00–5.20; ALE > PLE > PME > AME; palp 7.70 (2.87 + 1.13 + 1.68 + 2.02), leg I 9.57 (3.07 + 1.70 + 1.70 + 1.98 + 1.12), leg II 9.64 (2.95 + 1.68 + 1.61 + 2.08 + 1.32), leg III 9.60 (2.68 + 1.69 + 1.50 + 2.30 + 1.43), leg IV 14.21 (4.00 + 1.92 + 2.55 + 3.83 + 1.91).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium indistinct (Fig.
The species epithet, a noun in apposition, refers to the cryptic nature of this species discovery.
The species is endemic to the Japanese island Okinawajima (Fig.
Heptathela helios
Tanikawa & Miyashita, 2014: 68 (holotype: male (NSMT-Ar 12851), from Kunigami-son, Okinawajima, Japan, collected by A. Tanikawa on 26 May 2010, matured on 9 September 2012, deposited in
Males of H. helios can be distinguished from those of all other Okinawa group Heptathela species by the serrated contrategulum margin and the hooked tegular marginal apophysis, the ovate, indistinctly rugose conductor with a poorly serrated margin (Fig.
Male and female genital anatomy of Heptathela helio Tanikawa & Miyashita, 2014 A–G 2440A (short for XUX-2012-440A) H, K 2432 I, L 2434 J, M 2437 A palp prolateral view B palp ventral view C palp retrolateral view D–G palp distal view H–J vulva dorsal view K–M vulva ventral view. Scale bar: 0.5 mm.
Male. Carapace and opisthosoma brown, with dark brown tergites; cheliceral groove with 14 denticles; tergites closed to each other; seven spinnerets. Measurements: BL 9.88, CL 5.30, CW 4.81, OL 4.58, OW 2.92; ALE > PLE > PME > AME; leg I 16.56 (4.55 + 1.90 + 3.48 + 4.33 + 2.30), leg II 17.19 (4.50 + 2.00 + 3.34 + 4.68 + 2.67), leg III 18.35 (4.35 + 1.97 + 3.40 + 5.53 + 3.10), leg IV 23.37 (5.52 + 2.10 + 4.47 + 7.48 + 3.80).
Palp. The bulb of the two male specimens relatively distorted. Prolateral side of paracymbium unpigmented and unsclerotised, numerous setae and spines at the tip of paracymbium (Fig.
Females (N = 7). Carapace and opisthosoma colour as in male; cheliceral groove with 12–14 pronounced denticles; opisthosoma with 12 well-separated tergites; seven spinnerets. Measurements: BL 11.70–14.45, CL 5.29–6.70, CW 4.29–5.81, OL 6.54–8.18, OW 4.70–6.63; ALE > PLE > PME > AME; palp 12.29 (4.20 + 2.03 + 2.63 + 3.43), leg I 14.31 (4.51 + 2.40 + 2.65 + 3.08 + 1.67), leg II 14.04 (4.25 + 2.35 + 2.53+ 3.11 + 1.80), leg III 13.75 (3.92 + 2.40 + 2.48 + 2.67 + 2.28), leg IV 21.77 (5.90 + 2.88 + 3.75 + 6.11 + 3.13).
Female genitalia. A pair of depressions on the ventro-lateral part of genital atrium (Fig.
We identified the specimens collected from Ginama Dam, Okinawa, as H. helios based on evidence from morphology and COI barcode genetic distance compared with the male holotype and paratype (NSMT-Ar 12851, NSMT-Ar 12855) of H. helios in
JAPAN · 2 ♂♂, 8 ♀♀; Okinawa-ken, Kunigami-son, Ginama Dam; 26.84N, 128.26E; alt. 150 m; 24 December 2012; D. Li, F.X. Liu and X. Xu leg.; XUX-2012-432 to 440C.
The species is endemic to the Japanese island Okinawajima (Fig.
We thank Zoltán Korsós, Mamoru Toda, and Bo Wu for assistance in the field, and the staff of the Centre for Behavioural Ecology and Evolution (CBEE, Hubei University) for all their help and support throughout this study. We acknowledge constructive and insightful comments on the manuscript from Ingi Agnarsson, Jason Bond, Rebecca Godwin, and Xiang Xu. We thank Gonzalo Giribet, Laura Leibensperger, Stefan Friedrich, Danilo Harms, and Jason A. Dunlop for information on the types and for facilitating loans. This study was supported in part by the grants from National Natural Sciences Foundation of China (NSFC-31601850; NSFC-31272324), the Hunan Provincial Natural Science Foundation of China (2017JJ3202), Singapore Ministry of Education AcRF Tier 1 grant (R-154-000-A52-114) to DL, the Japan Society of Promotion of Science (JSPS-21540487), the Slovenian Research Agency (P1-10236 and J1-6729), and the bilateral inter-governmental S&T exchange project between China and Slovenia (12-8).
Table S1
Data type: species data