Monograph |
Corresponding author: Shuqiang Li ( lisq@ioz.ac.cn ) Academic editor: Yuri Marusik
© 2016 Yucheng Lin, Francesco Ballarin, Shuqiang 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:
Lin Y, Ballarin F, Li S (2016) A survey of the spider family Nesticidae (Arachnida, Araneae) in Asia and Madagascar, with the description of forty-three new species. ZooKeys 627: 1-168. https://doi.org/10.3897/zookeys.627.8629
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Forty-three new species of Nesticidae are described from China, Indonesia, Philippines, Singapore, Thailand, Vietnam and Madagascar, and two new junior synonyms are suggested. A new genus, Speleoticus gen. n., is described with Nesticus navicellatus Liu & Li, 2013 as the type species, and four species are transferfed from Nesticus, i.e., Speleoticus globosus (Liu & Li, 2013), comb. n., S. libo (Chen & Zhu, 2005), comb. n., S. navicellatus (Liu & Li, 2015), comb. n. and S. uenoi (Yaginuma, 1972), comb. n. The new species described in this paper belong to four genera and are: Hamus cornutus sp. n. (♂♀), H. kangdingensis sp. n. (♂), H. luzon sp. n. (♀), H. mangunensis sp. n. (♂), Nescina kohi sp. n. (♂♀), Nesticella baiseensis sp. n. (♂♀), N. baobab sp. n. (♂), N. caeca sp. n. (♂♀), N. chongqing sp. n. (♀), N. dazhuangensis sp. n. (♂♀), N. fuliangensis sp. n. (♂♀), N. gazuida sp. n. (♀), N. gongshanensis sp. n. (♀), N. griswoldi sp. n. (♂♀), N. hongheensis sp. n. (♂♀), N. huomachongensis sp. n. (♂♀), N. jingpo sp. n. (♀), N. kaohsiungensis sp. n. (♂♀), N. lisu sp. n. (♂♀), N. liuzhaiensis sp. n. (♀), N. nandanensis sp. n. (♂♀), N. phami sp. n. (♂♀), N. potala sp. n. (♀), N. qiaoqiensis sp. n. (♀), N. qiongensis sp. n. (♂♀), N. robusta sp. n. (♂♀), N. rongtangensis sp. n. (♂), N. sanchaheensis sp. n. (♂♀), N. sulawesi sp. n. (♀), N. sumatrana sp. n. (♂), N. tibetana sp. n. (♂♀), N. vanlang sp. n. (♀), N. wanzaiensis sp. n. (♂♀), N. xiongmao sp. n. (♂♀), N. xixia sp. n. (♂♀), N. yanbeiensis sp. n. (♂♀), N. yao sp. n. (♀), N. zhiyuani sp. n. (♂♀), Pseudonesticus dafangensis sp. n. (♂♀), P. miao sp. n. (♂♀), P. spinosus sp. n. (♂♀), P. wumengensis sp. n. (♀), P. ziyunensis sp. n. (♂♀). Nesticella inthanoni (Lehtinen & Saaristo, 1980), syn. n. is synonymised with N. mollicula(Thorell, 1898); N. taiwan Tso & Yoshida, 2000, syn. n. is synonymised with N. odonta (Chen, 1984). The female of Nesticella connectens Wunderlich, 1995, so far unknown, is described and recorded from Thailand. Nesticidae are reported from Madagascar for the first time. Nesticella nepalensis (Hubert, 1973) is recorded for the first time from China. Types of Nesticella odonta (Chen, 1984), N. songi Chen & Zhu, 2004 and N. yui Wunderlich & Song, 1995 are re-examined and photographed. The entire genus Nesticella is reviewed, and four species groups are recognised. DNA barcodes of the new species are obtained to confirm their correct identifications.
Nesticids, taxonomy, new taxa, cave, leaf litter
Nesticidae is a small family of spiders with an almost worldwide distribution, being absent only in Siberia, Central Asia, Northern and Southern Africa and at high latitudes. So far, 233 extant species belonging to 13 genera (
Traditionally the family Nesticidae has been considered rather uniform since established by Simon in 1894, with only one or very few genera included. Only in 1980 Lehtinen & Saaristo made a preliminary worldwide revision, grouping them in 10 genera and two tribes, Nesticini and Nesticellini, with the exception of the Nearctic and Neotropical genera Eidmannella Roewer, 1935 and Gaucelmus Keyserling, 1884 which were considered separately. The tribe Nesticini included the “long-legged” species restricted to caves and consisted of the genera Nesticus Thorell, 1869 from Europe and the Americas, Cyclocarcina Komatsu, 1942 endemic to Japan and Carpathonesticus Lehtinen & Saaristo, 1980 and Typhlonesticus Kulczyński, 1914 distributed in the Carpathian Mountains, Caucasus and around the Mediterranean Basin. The tribe Nesticellini contained all the so called “short-legged” nesticids distributed in the Oriental-Australian region that mostly live in leaf litter of tropical forests. According to the authors, this tribe consists of two main genera, Howaia Lehtinen & Saaristo, 1980 and Nesticella Lehtinen & Saaristo, 1980 separated by palpal morphological characters; these genera were subsequently synonymized by
Since the work of
In the last 15 years several collecting trips were carried out by Chinese, American and European arachnologists in Madagascar, China and Southeast Asia including Laos, Myanmar, Vietnam, Thailand, Singapore, Indonesia and Philippines. This effort allowed the collection many spiders, including numerous specimens of the family Nesticidae and was followed by the preliminary description of some new species (
The main aim of this paper is to report forty-three new species belonging to the genera Hamus, Nescina, Nesticella and Pseudonesticus. The new genus, Speleoticus gen. n., which includes the Japanese species S. uenoi (Yaginuma, 1972), and the remaining Chinese species previously placed in Nesticus, is established based on palpal and epigynal morphology.
Specimens used in this study were collected by hand or sifting leaf litter in China, Vietnam, Thailand, Philippines, Indonesia and Madagascar and immediately preserved in a 95% ethanol solution. All samples were examined using a Leica M205 C stereomicroscope and photographed with an Olympus c7070 wide zoom digital camera (7.1 megapixels). Images were mounted using Helicon Focus 3.10.3 software (
A partial fragment (625 bp) of the mitochondrial gene cytochrome c oxidase subunit I (COI) was amplified and sequenced to obtain the genetic distances between morphologically similar species and to confirm identifications and the sex pairing accuracy. Additionally, sequences of Hamus bowoensis Ballarin & Li, 2015, Nescina minuta Ballarin & Li, 2015, Nesticella connectens Wunderlich, 1995 and Nesticella odonta (Chen, 1984) were included; however, we were unable to obtain good extractions from Hamus kangdingensis sp. n., H. mangunensis sp. n., Nesticella lisu sp. n., N. xiongmao sp. n. and N. rongtangensis sp. n., and these were consequently excluded.
The primers used are: LCO1490 (5’-GGTCAACAAATCATCATAAAGATATTGG-3’) and CHR2 (5’-GGATGGCCAAAAAATCAAAATAAATG-3’). Raw sequences were edited and assembled using BioEdit v.7.2.5 (
Species | GenBank accession number | Collection localities |
---|---|---|
Hamus bowoensis Ballarin & Li, 2015 | KX866931 | China, Tibet, Bowo Co., Kaduo Village |
Hamus cornutus sp. n. | KX866932 | China, Guangxi, Pingxiang Co., Sanzhishan Cave |
Hamus luzon sp. n. | KX866933 | Philippines, Luzon Island, Mountain, Bontoc Town, rain forest |
Nescina kohi sp. n. | KX866934 | Singapore, Central Catchment Nature Reserve |
Nescina minuta Ballarin & Li, 2015 | KX866935 | China, Yunnan, Xishuangbanna, Mengla Co., Xiaolongha Village, Gougu forest |
Nesticella baiseensis sp. n. | KX866936 | China, Guangxi, Lingyun Co., Shuiyuan Cave |
Nesticella baobab sp. n. | KX866937 | Madagascar, Fianarantsoa, Parc National Ranomafana, Vohiparara |
Nesticella caeca sp. n. | KX866938 | China, Guizhou, Tianzhu Co., Liuhe Village, Jinshan Cave |
Nesticella chongqing sp. n. | KX866939 | China, Chongqing City, Beibei Dist., Dajiang Village, Xiaofang Cave |
Nesticella connectens Wunderlich, 1995 | KX866940 | Thailand, Satun, Trang Dist., Beating Cave |
Nesticella dazhuangensis sp. n. | KX866941 | China, Yunnan, Tengchong Co., Dazhuang Village |
Nesticella fuliangensis sp. n. | KX866942 | China, Jiangxi, Fuliang Co., Zhuxian Cave |
Nesticella gazuida sp. n. | KX866943 | China, Guizhou, Kaili City, Gazuida Cave |
Nesticella gongshanensis sp. n. | KX866944 | China, Yunnan, Gongshan Co., Langdang Village |
Nesticella griswoldi sp. n. | KX866945 | Madagascar, Toliara, Forest Classee Tsitongambarika, Cascade hiking trails |
Nesticella hongheensis sp. n. | KX866946 | China, Yunnan, Pingbian Co. |
Nesticella huomachongensis sp. n. | KX866947 | China, Hunan Province, Huaihua City, Chenxi Co., Huomachong Town, Yanzi Cave |
Nesticella jingpo sp. n. | KX866948 | China, Yunnan, Tengchong Co., Gaoligongshan Mountain National Park |
Nesticella kaohsiungensis sp. n. | KX866949 | China, Taiwan, Nantou Co., Hui Sun |
Nesticella liuzhaiensis sp. n. | KX866950 | China, Guangxi, Nandan Co., Longli Village, near the Dixia River |
Nesticella nandanensis sp. n. | KX866951 | China, Guangxi, Nandan Co., Encun Village, Encun Cave |
Nesticella odonta (Chen, 1984) | KX866952 | China, Jiangxi, Shangli Co., Zhanshan Village, Xiongxin Cave |
Nesticella phami sp. n. | KX866953 | Vietnam, Quang Ninh, Phong Nha Ke bang National Park, Sung Sot Cave |
Nesticella potala sp. n. | KX866954 | China, Tibet, Gyirong Co., Zalong Village |
Nesticella qiaoqiensis sp. n. | KX866955 | China, Sichuan, Baoxing Co., Zeyin Village |
Nesticella qiongensis sp. n. | KX866956 | China, Hainan, Diaoluoshan Mountain National Nature Reserve, Diaoluoshan Holiday Village |
Nesticella robusta sp. n. | KX866957 | China, Hunan, Yuanling Co., Qixian Cave |
Nesticella sanchaheensis sp. n. | KX866958 | China, Guizhou, Libo Co., Sanchahe Village, Sanchahe Cave |
Nesticella sulawesi sp. n. | KX866959 | Indonesia, South Sulawesi, Maros City, Cenrana Village |
Nesticella sumatrana sp. n. | KX866960 | Indonesia, West Sumatra, Payakumbuh City, Koto Tiggi Village, a cave without name, close to Imam Bonjol Cave |
Nesticella tibetana sp. n. | KX866961 | China, Tibet, Bowo Co., Yigong Town |
Nesticella vanlang sp. n. | KX866962 | Vietnam, Ninh Thuan, Nui Chua National Park |
Nesticella wanzaiensis sp. n. | KX866963 | China, Jiangxi, Wanzai Co., Dongkou Village, Zhushan Cave |
Nesticella xixia sp. n. | KX866964 | China, Henan, Xixia Co., Baihe Village, Yunhuabianfu Cave |
Nesticella yanbeiensis sp. n. | KX866965 | China, Guangxi, Lingchuan Co., Yanbei Village, Yanbei Cave |
Nesticella yao sp. n. | KX866966 | China, Guangxi, Gongcheng Co., Songlin Village, Houyan Cave |
Nesticella zhiyuani sp. n. | KX866967 | Indonesia, West Sumatra, Payakumbuh City, Koto Tiggi Village, a cave without name, close to Imam Bonjol Cave |
Pseudonesticus dafangensis sp. n. | KX866970 | China, Guizhou, Dafang Co., Sanhe Village, Yelaoda Cave |
Pseudonesticus miao sp. n. | KX866968 | China, Guizhou, Anshun City, Xixiu Dist., Xiaguantun Village, Duofan Cave |
Pseudonesticus spinosus sp. n. | KX866969 | China, Guizhou, Suiyang Co., Guihua Village, Mahuang Cave |
Pseudonesticus wumengensis sp. n. | KX866971 | China, Guizhou, Hezhang Co., Gaoyan Village, Tanjiayan Cave |
Pseudonesticus ziyunensis sp. n. | KX866972 | China, Guizhou, Ziyun Co., Mt. Wufeng, Wufeng Cave |
Abbreviations used in the text or figures are given in Table
Male palp | |
Ac | apex of the conductor |
C | conductor |
Cp (I-III) | processes of the conductor (I-III) |
Da | dorsal apophysis of the paracymbium |
Dp (I-II) | distal processes of the paracymbium (I-II) |
E | embolus |
Es | embolic spur |
Lf | lateral cymbial furrow |
Tg (I-II) | tegular apophysis (I-II) |
P | paracymbium |
St | subtegulum |
T | tegulum |
Ta (I-II) | terminal apophysis (I-II) |
Va (I-III) | ventral apophysis of the paracymbium (I-III) |
Epigyne | |
Co | copulatory opening |
Ep | epigynal plate |
Cd | copulatory duct |
Fd | fertilization duct |
S | spermatheca |
Sp | scape |
Vp | vulval pocket |
Ocular area | |
AER | anterior eye row |
ALE | anterior lateral eye |
AME | anterior median eye |
MOA | median ocular area |
PER | posterior eye row |
PLE | posterior lateral eye |
PME | posterior median eye |
Institutions | |
CASC | California Academy of Sciences, San Francisco, USA |
|
Institute of Zoology, Chinese Academy of Sciences, Beijing, China |
IBPN | Institute for Biological Problems of the North of the Russian Academy of Sciences, Magadan, Russia |
LKCNHM | Lee Kong Chian Natural History Museum, National University of Singapore |
MHBU | Museum of Hebei University, Baoding, China |
Hamus Ballarin & Li, 2015: 180.
Hamus bowoensis Ballarin & Li, 2015 from Tibet, China.
Hamus is easily distinguished from all the other Nesticidae, with the exception of Nescina, by the following combination of characters: for the males, a short paracymbium ending in only a tiny, hooked process (Dp) (usually more developed and ramificated in most other Nesticidae), a remarkable, almost round and ventrally flat bulb, a long, laminar conductor (C) with a counterclockwise course around the bulb (both absent in all the other genera except Nescina) and a compact and hook-like process of the conductor (Cp-I). Females are distinguished by the considerably large, almost round spermathecae (S), smaller in other genera. Males of the genus Hamus are separated from those of Nescina by the larger size, the larger terminal apophysis II (Ta-II) and the massive hook-like process I of the conductor (Cp-I) (Cp-I is reduced and flat in Nescina). Females can be separated by the wider spermathecae (S) and the shorter and simpler fertilization ducts (Fd).
Total length: 1.48–1.87 (male), 2.00–2.15 (female). Carapace rounded in males, more ovate in females, yellow or pale yellow. Cephalic area slightly raised, with sparse setae. Eyes ALE>PME=PLE>AME. Cervical groove and fovea indistinct. Chelicera with three promarginal teeth and multiple retromarginal tiny denticles on the fang furrow. Legs uniformly yellow; in male of H. bowoensis a tuft of hairs present on the prolateral margin of femur I. Opisthosoma grey or light yellow as the carapace, with long setae.
Male palp: tibia short, wider than long. Cymbium wide with well-developed lateral furrow about 2/3 of the cymbial length. Paracymbium short, compact and simple, weakly sclerotized, with a tiny, hooked distal process, a lobed ventral apophysis and a flat, translucent dorsal apophysis. Bulb almost round and ventrally flat. Terminal apophysis well-developed and strongly sclerotized with one or two processes, process I elongate or reduced, sometimes absent, process II always present and hook shaped. Tegular apophysis absent. Conductor long and laminar, starting at the prolateral margin of the bulb and following the embolus on the prolateral side with a counterclockwise course. Two strongly sclerotized processes located at the base of the conductor; the first short and compact, hook-like, the other elongate leading the terminal part of the embolus to the center of the bulb. Embolus long and slender starting from the retrolateral side of the bulb and bordering the tegulum with a semicircular clockwise course.
Epigyne: posterior margin of the epigynal plate weakly sclerotized and straight. Copulatory openings near the epigynal posterior margin. Spermathecae and fertilization ducts visible through the transparent tegument. Fertilization ducts relatively long and simple, copulatory ducts short, ventrally oriented. Spermathecae close to each other, wide and almost round or pyriform.
Hamus bowoensis Ballarin & Li, 2015, H. cornutus sp. n., H. kangdingensis sp. n., H. luzon sp. n., and H. mangunensis sp. n.
China (Guangxi, Sichuan, Tibet, Yunnan), Laos, Philippines (Luzon Island).
Hamus bowoensis Ballarin & Li, 2015: 180–183, figs 1–2 (♂♀).
Males can be separated from those of Hamus cornutus sp. n. and H. kangdingensis sp. n. by the tiny distal process of the paracymbium (Dp) (thicker in the other two species), by the presence of a well-developed terminal apophysis process I (Ta-I) and by the thicker apophysis process II with a less sharp point (Ta-II) (see
See
Forest leaf litter.
China (Tibet).
Holotype ♂ and paratypes 3♂3♀ (
The specific name derives from the Latin word “cornutus” = horned, and refers to the horn-like distal process of the paracymbium; adjective.
Males can be distinguished from those of Hamus bowoensis, H. mangunensis sp. n. and H. kangdingensis sp. n. by the absence of the terminal apophysis process I (Ta-I) (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.56. Carapace 0.80 long, 0.73 wide. Opisthosoma 0.77 long, 0.69 wide. Clypeus height 0.15. Sternum 0.47 long, 0.47 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.02. Carapace 0.90 long, 0.78 wide. Opisthosoma 1.22 long, 0.87 wide. Clypeus height 0.18. Sternum 0.56 long, 0.49 wide. Leg measurements: see Appendix
Forest leaf litter, cave.
China (Guangxi), Laos (Fig.
Holotype ♂ and paratype 1♂ (
The specific name is derived from the type locality; adjective.
Males can be easily distinguished from those of the other congeners by the caniniform distal process of the paracymbium (Dp) (Fig.
Habitus as in Fig.
Male palp (Fig.
Male (holotype). Total length 1.48. Carapace 0.79 long, 0.68 wide. Opisthosoma 0.78 long, 0.60 wide. Clypeus height 0.16. Sternum 0.48 long, 0.48 long. Leg measurements: see Appendix
Female. Unknown.
Forest leaf litter.
Known only from the type locality (Fig.
Holotype ♀ and paratype 1♀ (
The specific name is derived from the type locality, the island of Luzon in the Philippines; noun in apposition.
Females can be distinguished from those of Hamus bowoensis and H. cornutus sp. n. by the almost perfectly round spermathecae (S) and the shorter, narrower fertilization ducts (Fd) (Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.10. Carapace 0.95 long, 0.80 wide. Opisthosoma 1.34 long, 0.94 wide. Clypeus height 0.20. Sternum 0.57 long, 0.55 wide. Leg measurements: see Appendix
Male. Unknown.
Rainforest leaf litter.
Known only from the type locality (Fig.
Holotype ♂ (
The specific name is derived from the type locality, Mangun Village; adjective.
Hamus mangunensis sp. n. is easily distinguished from the other Hamus species by a tiny dentiform distal process of the paracymbium (Dp) (Fig.
Habitus cannot be properly described due to the very poor condition of the sample.
Male palp (Fig.
Female. Unknown.
Forest leaf litter.
Known only from the type locality (Fig.
Nescina Ballarin & Li, 2015: 183.
Nescina minuta Ballarin & Li, 2015 from Yunnan, China.
Males of Nescina are distinguished from those of the other genera of Nesticidae with the exception of Hamus and Wraios by a short and very simple paracymbium having only a lobed ventral apophysis (Va) and a sclerotized, hook-shaped distal process (Dp), by the almost round, ventrally flat bulb and by the elongate, laminar conductor (C) with only one small process (Cp-I). Males of Nescina can be separated from those of Hamus by the smaller terminal apophysis II (Ta-II) and the reduced, laminar process I of the conductor (Cp-I), and from those of Wraios by the shape of embolus. Diagnostic characters for the females are the long and twisted fertilization ducts (Fd) which have no similarities in the other genera. Furthermore Nescina is easily distinguished by the remarkably small size of the adults (total length less than 1.60) and the protruding cephalic area clearly separated from the thoracic area by a distinct cervical groove.
Total length: 1.30–1.56 (male), 1.44–1.57 (female). Carapace ovate, pale yellow. Cephalic area strongly elevated and clearly differentiated from the rest of the carapace. Eyes ALE=PLE>PME>AME, AER and PER procurved, MOA square. Cervical groove and fovea indistinct. Chelicera with six promarginal teeth and two retromarginal tiny denticles on the fang furrow. Sternum yellowish, heart-shaped. Opisthosoma yellowish, usually with four darker marks more or less visible. Spinnerets and colulus pale yellow.
Male palp: tibia short, as long as wide, with two retrodorsal trichobothria and several long setae. Cymbium almost round with long setae and long furrow on the retrolateral margin. Paracymbium simple, with a wide, lobed ventral apophysis and a hook-like distal process bent ventrally. Bulb ventrally flat. Terminal apophysis small and flat with two processes, Ta-I laminar, Ta-II spur-like. Tegular apophysis missing. Conductor laminar and long, starting from the anterior margin of the tegulum and following the embolus on the prolateral side with a counterclockwise course. One laminar process (Cp-I) at the subapical margin. Embolus filamentous starting from the retrolateral side of the bulb and continuing bordering the tegulum with a semicircular clockwise course.
Epigyne: wide, copulatory openings located near the epigynal posterior margin. Spermathecae and fertilization ducts can be observed through the transparent tegument. Spermathecae wide and globular, fertilization ducts long and twisted, laterally oriented. Copulatory ducts short.
Nescina minuta Ballarin & Li, 2015 and N. kohi sp. n.
China (Yunnan), Singapore (Fig.
Nescina minuta Ballarin & Li, 2015: 183, 188, figs 3–4 (♂♀).
Males can be separated from those of Nescina kohi sp. n. by the thicker terminal apophysis (Ta), by the different shape of the process I of the conductor (Cp-I) (see
Nescina kohi sp. n., holotype (male) and paratype (female). A Male habitus, dorsal view B Ditto, ventral view C Female habitus, dorsal view D Ditto, ventral view E Epigyne, ventral view F Vulva, ventral view G Vulva, dorsal view. Scale bars: A–D = 0.50 mm; E–G = 0.10 mm. Arrows indicate cervical groove in A and C.
See
Forest leaf litter.
China (Yunnan).
Holotype ♂ (LKCNHM), paratypes 2♀ (LKCNHM) and 1♂1♀ (
The new species is named after Joseph K. H. Koh, a pioneer spider researcher in Singapore; noun (name) in genitive case.
Nescina kohi sp. n. morphologically is very similar to N. minuta (see Figs
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.30. Carapace 0.69 long, 0.62 wide. Opisthosoma 0.65 long, 0.50 wide. Clypeus height 0.13. Sternum 0.46 long, 0.43 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 1.44. Carapace 0.65 long, 0.58 wide. Opisthosoma 0.82 long, 0.64 wide. Clypeus height 0.09. Sternum 0.45 long, 0.41 wide. Leg measurements: see Appendix
Forest leaf litter.
Known only from the type locality (Fig.
The molecular analysis confirms that Nescina kohi sp. n. differs from the type species, N. minuta (see Appendix
Nesticella
Lehtinen & Saaristo, 1980: 55;
Nesticus nepalensis Hubert, 1973 from Nepal.
Nesticella belongs to the tribe Nesticellinisensu Lehtinen & Saaristo (1980). The characters are rather variable within the different species-groups of Nesticella, however males can be recognized by the presence of a complex paracymbium usually lacking any dorsal apophysis (a few species with only one), with a simple or bifurcate distal process and one or two ventral processes. By contrast, males of the other small-sized nesticid genera distributed in the Old World (Hamus, Nescina, Wraios) have a shorter and more compact paracymbium. The uncoiled embolus instead of a long and coiled embolus, easily separate Nesticella from the small Pseudonesticus species. Females of Nesticella are recognized by the short, rectangular or lobed and often protruding scape and by the general shape of the copulatory ducts with thick, compact copulatory ducts lacking deep convolutions and located just below the coiled, thin fertilization ducts. The above-mentioned characters are not similar to any other genus of Nesticidae.
Total length: 1.60–2.94 (male), 2.00–3.36 (female). Carapace almost round in males, ovate in females, yellow or pale yellowish, only rarely dark. Cephalic area slightly elevated with sparse setae. Eight eyes in two rows with exceptions in cave-adapted species, ALE>PLE=PME>AME. AER slightly procurved, PER straight MOA trapezoidal, narrower in the front. In troglobitic species (e.g. N. caeca sp. n., N. gazuida sp. n.) eyes strongly reduced with AME sometimes absent. Cervical groove and fovea usually distinct, sometimes indistinct. Chelicera with three promarginal teeth and multiple retromarginal tiny denticles. Opisthosoma with long setae usually with a yellowish or greyish background and pairs of dark spots partially merged each other on the dorsal side, in some species uniformly black or greyish. Colors and marks can be more or less visible and sometimes reduced to being faint.
Male palp: tibia short and swollen, wider than long, with three retrodorsal trichobothria and sparse long setae. Cymbium with dense and thick long setae dorsal-distally. Paracymbium compact with a stocky, laminar or bifurcate distal process and a ventral apophysis with one or two processes of different length. Dorsal apophysis absent in some species. Terminal apophysis blunt, crest-like or elongate and pointed. Tegular apophysis usually well-developed, missing in mogera and quelpartensis-groups. Tegulum with a small additional tegular apophysis (Tg-II) in the species belonging to the nepalensis-group. Conductor long and curved, basally wide, gradually narrower near the tip, sometimes distally twisted and always with a sclerotized, short beak-shaped process. Embolus long and filamentous, starting from the retroventral margin of the tegulum and coiling with a half loop until reaching the apex of the conductor.
Epigyne: weakly sclerotized showing the internal structures. Scape lobed or squared, sometimes strongly reduced, sometimes well-developed. Copulatory openings at the lateral corners of the scape, under the scape in the phami-group. Spermathecae ovoid or almost round, usually small. Fertilization ducts long and thin reaching the spermathecae with one to five coils. Copulatory ducts short and thick, ventrally oriented, sometimes bent in the middle and directed laterally.
Mostly Pantropical (Brazil; Central and East Africa; East, South and Southeast Asia; Indochina; Melanesia) with few species occurring in the Russian Far East. Nesticella mogera is also found in Azerbaijan, Fiji, Hawaii and Europe, probably introduced by human activities.
Although the morphological characters of Nesticella species may support the hypothesis of two possible distinct genera as claimed by
Two recently-described species from Laos and Myanmar, Nesticella foelixi Grall & Jäger, 2016 and N. michaliki Grall & Jäger, 2016, show peculiar palpal and epigynal characters which don’t allow their clear placement into any Nesticella species-group in this paper. Some characters, such as shape of the scape, the wide spermathecae and the flat distal process of the paracymbium, are shared with phami-group species; however the thick embolus, the general shape of the process of the conductor, the terminal apophysis and the paracymbium are not shared with this species group. It is possible that these new species represent separate monospecific species-groups. Nevertheless, in absence of further data, these species are temporarily considered as incertae sedis until further analyses have been conducted.
Group features. Males of this species-group are characterized by the following combination of characters: a ventral apophysis of the paracymbium with two parallel processes (Va-I, Va-II), the first longer than the second and usually ending with a sharp tip; a distal process of the paracymbium with two branches (Dp-I, Dp-II) more or less developed and in some cases heavily reduced (e.g. N. hongheensis sp. n.) or partially absent (e.g. N. dazhuangensis sp. n. and N. caeca sp. n.); a well-developed tegular apophysis (Tg) directed retrolaterally and a small process of the conductor. Females belonging to this species-group are characterized by a protruding lobed scape (Sp) with a rounded tip. The scape can be more rarely short and very wide (wider then long). These features allow a relatively easy separation from females of the mogera and nepalensis-groups which have a squarer or narrower scape. The smaller spermathecae (S) easily distinguish females of the brevipes-group from those of the phami-group whereas the bent copulatory ducts (Cd) further separate them from those of the mogera and quelpartensis-groups.
Composition. Nesticella arcuata Liu & Li, 2013; N. baiseensis sp. n.; N. brevipes (Yaginuma, 1970); N. buicongchieni (Lehtinen & Saaristo, 1980); N. caeca sp. n.; N. chongqing sp. n.; N. dazhuangensis sp. n.; N. falcata Liu & Li, 2013; N. gazuida sp. n.; N. gracilenta Liu & Li, 2013; N. hongheensis sp. n.; N. jingpo sp. n.; N. kerzhneri (Marusik, 1987); N. lisu sp. n.; N. liuzhaiensis sp. n.; N. machadoi (Hubert, 1971)?; N. mollicula (Thorell, 1898); N. nandanensis sp. n.; N. odonta (Chen, 1984); N. okinawaensis (Yaginuma, 1979)?; N. qiaoqiensis sp. n.; N. qiongensis sp. n.; N. robusta sp. n.; N. sanchaheensis sp. n.; N. semicircularis Liu & Li, 2013; N. shanlinensis Liu & Li, 2013; N. songi Chen & Zhu, 2004; N. verticalis Liu & Li, 2013; N. xiongmao sp. n.; N. xixia sp. n. and N. yao sp. n.
Remarks. The placement of Nesticella machadoi (Hubert, 1971) and N. okinawaensis (Yaginuma, 1979) in the brevipes-group is provisional due to the poor original illustrations and descriptions which do not allow a clear understanding the diagnostic characters. A direct and detailed examination of the type material is necessary to confirm their affinity with this species-group.
Holotype ♂ and paratypes 2♀ (
The specific name is derived from the type locality; adjective.
The new species is closely related to Nesticella nandanensis sp. n., N. songi and N. verticalis. It can be separated from N. nandanensis sp. n. by the longer tegular apophysis (Ta), the larger angle formed by the distal processes of the paracymbium (Dp-I, Dp-II) (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.97. Carapace 1.47 long, 1.31 wide. Opisthosoma 1.72 long, 1.27 wide. Clypeus height 0.25. Sternum 0.90 long, 0.83 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 3.28. Carapace 1.37 long, 1.19 wide. Opisthosoma 2.09 long, 1.72 wide. Clypeus height 0.25. Sternum 0.84 long, 0.79 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Nesticus brevipes Yaginuma, 1970: 386 figs 1–2, 9–10 (♂♀).
Howaia
brevipes
:
Nesticella
brevipes
:
Nesticella
brevipes
:
Howaia
brevipes
:
See the
Males of Nesticella brevipes are distinguished from those of all the other species belonging to the brevipes-group by the well-developed and protruding dorsal apophysis (missing or only slightly protruding in the other species) and by the single, thin and sharp distal process of the paracymbium (usually bifurcated or, if single, thinner or blunt in the other species of the brevipes-group) (see
See
Forest leaf litter, cave.
China (Zhejiang), Japan, Korea, Russia (Sakhalin, South Kuril Islands).
Holotype ♂ and paratypes 2♀ (
The epithet derives from the Latin word “caecus” = blind, referring to the vestigial eyes; adjective.
Males and females of Nesticella caeca sp. n. can be easily recognized from those of the other species belonging to the brevipes-group, with the exception of N. gazuida sp. n. (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.43. Carapace 1.19 long, 1.08 wide. Opisthosoma 1.30 long, 0.87 wide. Clypeus height 0.22. Sternum 0.74 long, 0.66 wdie. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.50. Carapace 1.14 long, 1.00 wide. Opisthosoma 1.66 long, 1.25 wide. Clypeus height 0.20. Sternum 0.69 long, 0.67 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
The molecular analysis shows that there are certain differences between Nesticella caeca sp. n. and the close related N. gazuida sp. n. (see Appendix
Holotype ♀ and paratype 1♀ (
The specific name is derived from the type locality, the city of Chongqing; noun in apposition.
The new species is closely related to Nesticella odonta (see Figs
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.75. Carapace 1.30 long, 1.13 wide. Opisthosoma 1.47 long, 1.41 wide. Clypeus height 0.23. Sternum 0.75 long, 0.72 wide. Leg measurements: see Appendix
Male. Unknown.
Cave.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 2♂2♀ (
The specific name is derived from the type locality; adjective.
Males can be distinguished from those of the other species belonging to the brevipes-group by the stocky, unbranched distal process (Dp) of the paracymbium (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.56. Carapace 1.38 long, 1.25 wide. Opisthosoma 1.28 long, 1.00 wide. Clypeus height 0.23. Sternum 0.78 long, 0.75 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.59. Carapace 1.25 long, 1.09 wide. Opisthosoma 1.56 long, 1.16 wide. Clypeus height 0.19. Sternum 0.75 long, 0.68 wide. Leg measurements: see Appendix
Forest leaf litter.
Known only from the type locality (Fig.
Holotype ♀ (
The specific name is derived from the name of the cave where the species was collected; noun in apposition.
Females of Nesticella gazuida sp. n. are easily distinguished from those of the other species belonging to the brevipes-group, with the exception of N. caeca sp. n. (Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.83. Carapace 1.17 long, 1.16 wide. Opisthosoma 1.75 long, 1.34 wide. Clypeus height 0.22. Sternum 0.68 long, 0.66 wide. Leg measurements: see Appendix
Male unknown.
Cave.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 1♂4♀ (
The specific name is derived from the type locality; adjective.
Nesticella hongheensis sp. n. is easily distinguished from the majority of the species belonging to the brevipes-group, with the exception of N. semicircularis and N. lisu sp. n., by having a wide, triangular dorsal apophysis (Da) (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.15. Carapace 1.13 long, 1.05 wide. Opisthosoma 1.05 long, 1.02 wide. Clypeus height 0.21. Sternum 0.66 long, 0.72 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.25. Carapace 1.18 long, 1.03 wide. Opisthosoma 1.28 long, 0.95 wide. Clypeus height 0.22. Sternum 0.72 long, 0.71 wide. Leg measurements: see Appendix
In humid and shaded areas, including anthropogenic habitats.
Known only from the type locality (Fig.
Holotype ♀ (
The species is named after the Jingpo people, an ethnic minority living in the Gaoligongshan region of Yunnan Province; noun in apposition.
This new species is closely related to N. lisu sp. n. (Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.18. Carapace 0.98 long, 0.89 wide. Opisthosoma 1.30 long, 1.08 wide. Clypeus height 0.17. Sternum 0.65 long, 0.62 wide. Leg measurements: see Appendix
Male. Unknown.
Forest leaf litter, under stone and rock.
Known only from the type locality (Fig.
Holotype ♂ and paratype 1♀ (
The species is named after the Lisu people, an ethnic minority living in Yunnan Province; noun in apposition.
Males of Nesticella lisu sp. n. can be easily distinguished from those of the other species belonging to the brevipes-group by the squared, dorso-ventrally flat distal process of the paracymbium (Dp) with serrated margin (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.14. Carapace 1.08 long, 0.97 wide. Opisthosoma 1.08 long, 1.00 wide. Clypeus height 0.18. Sternum 0.68 long, 0.62 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 3.03. Carapace 1.16 long, 1.00 wide. Opisthosoma 2.09 long, 1.43 wide. Clypeus height 0.21. Sternum 0.73 long, 0.68 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Holotype ♀ and paratype 1♀ (
The specific name is derived from the type locality; adjective.
The new species can be easily distinguished from all the others belonging to the brevipes-group by the very large and squared short scape, the flat posterior margin and the nearly straight copulatory ducts visible through the transparent tegument which give the epigyne a U-shape. Nesticella robusta sp. n. (Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.13. Carapace 1.00 long, 0.87 wide. Opisthosoma 1.36 long, 1.11 wide. Clypeus height 0.15. Sternum 0.64 long, 0.49 wide. Leg measurements: see Appendix
Male. Unknown.
Cave.
Known only from the type locality (Fig.
Erigone mollicula Thorell, 1898: 318 (♂♀).
Howaia inthanoni Lehtinen & Saaristo, 1980: 55, fig. 13 (♀), syn. n.
Nesticella
mollicula
:
The authors had the opportunity to see photos of the type material of Nesticella mollicula from Myanmar preserved in the collection of the Museo di Storia Naturale di Genova, Italy. The morphology of the species is comparable with N. inthanoni described from Thailand by
Myanmar, Thailand.
Holotype ♂ and paratypes 1♂4♀ (
The specific name is derived from the type locality; adjective.
Nesticella nandanensis is closely related to N. baiseensis sp. n., N. songi, and N. verticalis. It can be separated from N. baiseensis sp. n. by the shorter, stockier tegular apophysis (Tg), the smaller angle formed by the distal processes (Dp) of the paracymbium (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.56. Carapace 1.34 long, 1.25 wide. Opisthosoma 1.30 long, 1.06 wide. Clypeus height 0.21. Sternum 0.80 long, 0.76 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.91. Carapace 1.44 long, 1.30 wide. Opisthosoma 1.65 long, 1.28 wide. Clypeus height 0.23. Sternum 0.86 long, 0.84 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Nesticus odontus Chen, 1984: 34, figs 1–6 (♂♀).
Nesticella
odonta
:
Nesticus
odontus
:
Nesticella
odonta
:
Nesticella
odonta
:
Nesticella taiwan Tso & Yoshida, 2000: 13, figs 1–6 (♂), syn. n.
Paratypes of Nesticella odonta 1♂ and 2♀ (
This species can be easily recognized from the others belonging to the brevipes-group, with the exception of N. xiongmao sp. n. (Fig.
See
Forest leaf litter, cave.
South China (Fig.
The type material of Nesticella taiwan Tso & Yoshida, 2000 was not found in the Department of Biology at Tung Hai University, Taichung. However, the drawings of the male holotype by Tso & Yoshida clearly show the diagnostic characters of the species: the hook-like, pointed distal process I, the lobed distal process II, the general shape of the paracymbium when viewed ventrally and the pointed, sclerotized tegular apophysis. All these characters are compatible with the the palpal morphology of N. odonta from mainland China (compare Fig.
Holotype ♀ (
The specific name is derived from the type locality; adjective.
This new species is similar to Nesticella gracilenta (see
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.77. Carapace 1.09 long, 0.96 wide. Opisthosoma 1.76 long, 1.55 wide. Clypeus height 0.19. Sternum 0.58 long, 0.65 wide. Leg measurements: see Appendix
Male. Unknown.
Cave.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 2♀ (
The specific name derives from the Chinese pinyin for “Qiong”, and refers to an alias name of Hainan Island where the species was collected; adjective.
Nesticella qiongensis sp. n. can be distinguished from the majority of the species belonging to the brevipes-group, with the exception of N. falcata, N. liuzhaiensis sp. n. and N. robusta sp. n., by the absence of a dorsal apophysis (Da) (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.88. Carapace 1.04 long, 0.92 wide. Opisthosoma 0.84 long, 0.79 wide. Clypeus height 0.17. Sternum 0.60 long, 0.59 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.18. Carapace 1.09 long, 0.94 wide. Opisthosoma 1.16 long, 0.92 wide. Clypeus height 0.18. Sternum 0.63 long, 0.63 wide. Leg measurements: see Appendix
Forest leaf litter.
Known only from the type locality (Fig.
Holotype ♂ and paratype 1♀ (
The specific name is derived from the Latin word “robustus” = robust, sturdy, and refers to the strong distal process of the paracymbium in the male; adjective.
Males of Nesticella robusta sp. n. can be easily recognized by the single, thick and sturdy distal process of the paracymbium (Dp) with a beak-like shape and the very short ventral process II (Va-II) that has no similarities to any other species of the brevipes-group. The morphologically closest species is N. qiongensis sp. n. from which N. robusta sp. n. can be separated by the slimmer and sharper distal process of the paracymbium (Dp) and the shorter ventral process II (Va-II) (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.98. Carapace 1.14 long, 1.02 wide. Opisthosoma 0.90 long, 0.88 wide. Clypeus height 0.17. Sternum 0.61 long, 0.58 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.16. Carapace 1.07 long, 0.96 wide. Opisthosoma 1.15 long, 0.96 wide. Clypeus height 0.18. Sternum 0.60 long, 0.58 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 3♀ (
The specific name is derived from the type locality; adjective.
The new species is closely related to Nesticella gazuida sp. n. (see Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.78. Carapace 1.31 long, 1.18 wide. Opisthosoma 1.56 long, 1.15 wide. Clypeus height 0.23. Sternum 0.80 long, 0.78 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.81. Carapace 1.31 long, 1.16 wide. Opisthosoma 1.66 long, 1.38 wide. Clypeus height 0.22. Sternum 0.81 long, 0.75 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Nesticella songi Chen & Zhu, 2004: 87, figs 1–7 (♂♀).
Holotype ♂ and paratypes 1♂2♀ (MHBU), CHINA: Guizhou Province, Libo County, Maolan National Nature Reserve, Shuipa Village, Shuipashui Cave (25.40000°N, 107.80000°E), 7.VII.2001, H. Chen leg.
Nesticella songi is closely related to N. nandanensis sp. n., N. baiseensis sp. n. and N. yao sp. n. Males of the new species can be separated from those of N. nandanensis sp. n. by the blunter tip of the tegular apophysis (Tg) (Fig.
See Figs
Cave.
China (Guizhou) (Fig.
Holotype ♂ and paratypes 8♂5♀ (
The specific name derives from the Chinese pinyin “xióng māo” = panda, and refers to the type locality of the new spider species, located in the type locality of the giant panda; noun in apposition.
The new species can be easily distinguished from all the species belonging to the brevipes-group, with the exception of Nesticella odonta and N. chongqing sp. n., by the sharp, hook-like distal process I of the paracymbium (Dp-I) that is bent downward (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.45. Carapace 1.18 long, 1.09 wide. Opisthosoma 1.45 long, 1.06 wide. Clypeus height 0.23. Sternum 0.75 long, 0.70 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.91. Carapace 1.17 long, 0.99 wide. Opisthosoma 1.84 long, 1.64 wide. Clypeus height 0.23. Sternum 0.71 long, 0.66 wide. Leg measurements: see Appendix
Forest leaf litter.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 3♂5♀ (
The specific name is derived from the type locality; adjective.
This new species is closely related to Nesticella sanchaheensis sp. n. and N. semicircularis. It can be separated from the former species by the sharper tip of the tegular apophysis (Tg), the much thinner and sharp ventral process I (Va-I) and the smaller and narrower distal process I of the paracymbium (Dp-I) when observed in retrolateral view (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.84. Carapace 1.48 long, 1.33 wide. Opisthosoma 1.50 long, 1.06 wide. Clypeus height 0.26. Sternum 0.88 long, 0.82 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.97. Carapace 1.42 long, 1.25 wide. Opisthosoma 1.88 long, 1.56 wide. Clypeus height 0.27. Sternum 0.84 long, 0.80 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Holotype ♀ and paratype 1♀ (
The name derives from the Yao people, an ethnic minority living in the type locality; noun in apposition.
The new species is closely related to Nesticella songi Chen & Zhu, 2004 (see Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.84. Carapace 1.36 long, 1.14 wide. Opisthosoma 1.72 long, 1.13 wide. Clypeus height 0.22. Sternum 0.80 long, 0.78 wide. Leg measurements: see Appendix
Male. Unknown.
Cave.
Known only from the type locality (Fig.
Group features. Males belonging to this species-group are characterized by the following combination of characters: paracymbium having a single ventral apophysis (Va) with a sharp tip bent inward and a squared, stumpy single distal process (Dp); a terminal apophysis (Ta) that is usually flat, basally broad and with a coarse, granulate surface; tegular apophysis absent (Tg); a wide, strongly sclerotized, hook-like process of the conductor (Cp). Females are recognized by the straight and almost parallel copulatory ducts (Cd) and only slightly diverging distally. The massive squared scape (Sp) (wider, narrower or lobed in the other species-groups) that is always protruding far beyond the epigastric furrow further separates this group from the others.
Composition. Nesticella apiculata Liu & Li, 2013, N. fuliangensis sp. n., N. helenensis (Hubert, 1977), N. huomachongensis sp. n., N. mogera (Yaginuma, 1972), N. rongtangensis sp. n., N. wanzaiensis sp. n. and N. yanbeiensis sp. n.
Holotype ♂ and paratypes 2♂6♀ (
The specific name is derived from the type locality; adjective.
This new species is closely related to Nesticella huomachongensis sp. n. (Figs
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 3.16. Carapace 1.47 long, 1.28 wide. Opisthosoma 1.91 long, 1.34 wide. Clypeus height 0.23. Sternum 0.91 long, 0.80 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 3.56. Carapace 1.56 long, 1.31 wide. Opisthosoma 2.28 long, 1.69 wide. Clypeus height 0.22. Sternum 0.97 long, 0.81 wide. Leg measurements: See Appendix
Cave.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 2♀ (
The specific name is derived from the type locality; adjective.
Nesticella huomachongensis sp. n. is closely related to N. wanzaiensis sp. n. (see Figs
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.66. Carapace 1.38 long, 1.21 wide. Opisthosoma 1.44 long, 1.02 wide. Clypeus height 0.18. Sternum 0.71 long, 0.75 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 3.28. Carapace 1.44 long, 1.22 wide. Opisthosoma 1.88 long, 1.43 wide. Clypeus height 0.24. Sternum 0.81 long, 0.78 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Nesticus terrestris Yaginuma, 1970: 390, fig. 7 (♂, mismatched).
Nesticus mogera Yaginuma, 1972: 621, fig. 1 (♂♀).
Howaia
mogera
:
Howaia
mogera
:
Nesticella
mogera
:
See the World Spider Catalog for the full list of references.
Males of Nesticella mogera are recognized from those of the other species belonging to the mogera-group, with the exception of N. rongtangensis sp. n., by the very thin ventral apophysis and the blunter distal process of the paracymbium (see
See
A wide range of different habitats, like forest leaf litter, fields, caves, including greenhouses and city parks.
Azerbaijan, China, Korea, Japan, Hawaii, Fiji and Europe (introduced).
Holotype ♂ (
The specific name is derived from the type locality; adjective.
The lack of eyes is a clear, diagnostic character that allows the distinction of N. rongtangensis sp. n. from all other species of the mogera-group (Fig.
Habitus as in Fig.
Male palp (Fig.
Male (holotype). Total length 1.60. Carapace 0.92 long, 0.88 wide. Opisthosoma 0.70 long, 0.64 wide. Clypeus height 0.15. Sternum 0.62 long, 0.45 wide. Leg measurements: see Appendix
Female. Unknown.
Cave.
Known only from the type locality (Fig.
Holotype ♂ (
The specific name is derived from the type locality; adjective.
This new species is closely related to Nesticella huomachongensis sp. n. (see Figs
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.66. Carapace 1.27 long, 1.19 wide. Opisthosoma 1.50 long, 0.94 wide. Clypeus height 0.22. Sternum 0.83 long, 0.71 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 3.15. Carapace 1.34 long, 1.25 wide. Opisthosoma 1.82 long, 1.63 wide. Clypeus height 0.21. Sternum 0.76 long, 0.73 wide. Leg measurements: see Appendix
Cave.
Jiangxi, China (Fig.
Holotype ♂ and paratypes 3♀ (
The specific name is derived from the type locality; adjective.
Males can be separated from other species of the mogera-group by the relatively slim and wavy shape of the process of the conductor (Cp) and by the solid, rectangular distal process of the paracymbium (Dp) (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.94. Carapace 1.38 long, 1.25 wide. Opisthosoma 1.66 long, 1.17 wide. Clypeus height 0.21. Sternum 0.86 long, 0.79 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 3.36. Carapace 1.49 long, 1.31 wide. Opisthosoma 2.00 long, 1.41 wide. Clypeus height 0.22. Sternum 0.90 long, 0.82 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Group features. Males belonging to this species-group can be recognized by the following combination of characters: a ventral apophysis of the paracymbium with two lobed processes bent inward (Va-I, Va-II), Va-I longer and thicker than Va-II and usually ending with an almost round tip; a distal process of the paracymbium with one (more rarely) or two processes (Dp-I and Dp-II), Dp-I protruded ventrally, Dp-II retrolaterally protruded; a usually elongate terminal apophysis (Ta); a small, strongly sclerotized, tegular apophysis (Tg) protruding outward, an additional more or less developed tegular apophysis (Tg-II), and a sclerotized, hook-like process of the conductor (Cp). A diagnostic character for the females is the short or very short, narrow scape (Sp) (approximately as long as wide but remarkably narrower than the species of the mogera-group), which is barely visible in some species (wider, longer or with a rounded tip in the females of the other species-groups). Furthermore, the smaller size of the spermathecae (S) easily distinguish females of the nepalensis-group from those of the phami-group while the thick and bent copulatory ducts (Cd) separate them from those of the mogera and quelpartensis-groups
Composition. Nesticella aelleni (Brignoli, 1972), N. africana (Hubert, 1970), N. baobab sp. n., N. beccus Grall & Yäger, 2016, N. benoiti (Hubert, 1970), N. chillagoensis Wunderlich, 1995, N. connectens Wunderlich, 1995, N. ducke Rodrigues & Buckup, 2007, N. gongshanensis sp. n., N. griswoldi sp. n., N. laotica Grall & Yäger, 2016, N. murici Rodrigues & Buckup, 2007, N. nepalensis (Hubert, 1973), N. potala sp. n., N. proszynkii (Lehtinen & Saaristo, 1980), N. renata (Bourne, 1980), N. robinsoni Lehtinen & Saaristo, 1980, N. sechellana (Simon, 1898), N. sogi Lehtinen & Saaristo, 1980, N. sulawesi sp. n., N. utuensis (Bourne, 1980), N. taurama Lehtinen & Saaristo, 1980, N. tibetana sp. n., N. vanlang sp. n., N. yui Wunderlich & Song, 1995, and N. zhiyuani sp. n.
Holotype ♂ and paratype 1♂ (CASC), MADAGASCAR: Fianarantsoa Province, Parc National Ranomafana, Vohiparara, 3.6 km West of Ranomafana, primary montane rain forest (21.23738°S, 47.39737°E, 1150 m), 13–14.I.2009, C. Griswold, A. Saucedo & H. Wood leg.
The specific name is derived from from the word “baobab”, the common name for trees of the genus Adansonia which are common in Madagascar; noun in apposition.
The new species is similar to Nesticella benoiti (see
Habitus as in Fig.
Male palp (Fig.
Male (holotype). Total length 1.96. Carapace 1.03 long, 0.96 wide. Opisthosoma 1.02 long, 0.80 wide. Clypeus height 0.18. Sternum 0.60 long, 0.57 wide. Leg measurements: see Appendix
Female. Unknown.
Rain forest leaf litter.
Known only from the type locality (Fig.
Nesticella connectens Wunderlich, 1995: 568, figs 40–42 (♂).
1♂2♀ (
This species is similar to Nesticella tibetana sp. n. (see Figs
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male. Total length 1.93. Carapace 1.05 long, 0.98 wide. Opisthosoma 1.00 long, 0.76 wide. Clypeus height 0.19. Sternum 0.65 long, 0.63 wide. Leg measurements: see Appendix
Female. Total length 2.00. Carapace 1.06 long, 0.90 wide. Opisthosoma 1.18 long, 0.90 wide. Clypeus height 0.18. Sternum 0.65 long, 0.59 wide. Leg measurements: see Appendix
Forest leaf litter, cave.
Malaysia, Thailand (first record) (Fig.
Nesticella connectens was described by
Holotype ♀ (
The specific name is derived from the type locality; adjective.
This new species is closely related to Nesticella yui (see Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.38. Carapace 1.24 long, 1.07 wide. Opisthosoma 1.22 long, 1.10 wide. Clypeus height 0.23. Sternum 0.73 long, 0.66 wide. Leg measurements: see Appendix
Male. Unknown.
Forest leaf litter.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 1♂1♀ (CASC), MADAGASCAR: Toliara Province, Forest Classee Tsitongambarika, Cascade hiking trail, 7.5 km Northwest of Taolagnaro, primary montane rain forest (24.98664°S, 46.92631°E, 100 m), 24.XII.2008, F. Alvarez-Padilla & H. Wood leg.
The new species is named after Dr. Charles Griswold, a leading spider taxonomist from the USA; noun (name) in genitive case.
The new species is similar to Nesticella benoiti (see
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.93. Carapace 1.05 long, 0.98 wide. Opisthosoma 1.00 long, 0.76 wide. Clypeus height 0.19. Sternum 0.65 long, 0.63 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.00. Carapace 1.06 long, 0.90 wide. Opisthosoma 1.18 long, 0.90 wide. Clypeus height 0.18. Sternum 0.65 long, 0.59 wide. Leg measurements: see Appendix
Rain forest leaf litter.
Known only from the type locality (Fig.
Nesticus nepalensis Hubert, 1973: 165, figs 2–5 (♂♀).
Nesticella
nepalensis
:
Nesticella
nepalensis
:
1♂ (IBPN), INDIA: Uttarakhand State, Gobind Ghat Village (30.625°N, 79.55833°E, 1900 m), 17–23.V.1999, Y.M. Marusik leg. 2♀ (
Nesticella nepalensis is similar to N. connectens (see Figs
See Figs
Forest leaf litter, under stone.
China (Tibet) (first record) (Fig.
Holotype ♀ (
The specific name derives from the Potala Palace, one of the most famous Buddhist holy land in the world, which is located near the type locality of the species; noun in apposition.
The new species is closely related to Nesticella tibetana sp. n. (see Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 3.20. Carapace 1.37 long, 1.20 wide. Opisthosoma 2.13 long, 2.18 wide. Clypeus height 0.25. Sternum 0.84 long, 0.83 wide. Leg measurements: see Appendix
Male. Unknown.
Forest leaf litter.
Known only from the type locality (Fig.
Holotype ♀ and paratypes 2♀ (
The specific name is derived from the Island of Sulawesi were this species was collected; noun in apposition.
This new species can be distinguished from all the others of the nepalensis-group with the exception of Nesticella yui by the small spermathecae (S) and the twisted copulatory ducts (Cd) firstly bent inward and then outward before reaching the spermathecae (Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.33. Carapace 1.18 long, 1.05 wide. Opisthosoma 1.15 long, 1.00 wide. Clypeus height 0.19. Sternum 0.71 long, 0.68 wide. Leg measurements: see Appendix
Male. Unknown.
Rain forest leaf litter.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 1♂1♀ (
The specific name is derived from the type locality; adjective.
The new species is closely related to Nesticella connectens (see Figs
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.08. Carapace 1.09 long, 1.00 wide. Opisthosoma 1.00 long, 0.98 wide. Clypeus height 0.20. Sternum 0.67 long, 0.67 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.20. Carapace 1.10 long, 1.01 wide. Opisthosoma 1.20 long, 1.04 wide. Clypeus height 0.20. Sternum 0.66 long, 0.66 wide. Leg measurements: see Appendix
Forest leaf litter.
Known only from the type locality (Fig.
Holotype ♀ (
The specific name is derived from Van Lang, an ancient name for Vietnam; noun in apposition.
This new species is distinguished from all the other species of the nepalensis-group with the exception of Nesticella sogi by the straight fertilization ducts (Fd) and the wide spermathecae (S) which are close to each other, separated by less than a half of their diameter (Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.23. Carapace 1.11 long, 0.96 wide. Opisthosoma 1.45 long, 1.55 wide. Clypeus height 0.17. Sternum 0.61 long, 0.65 wide. Leg measurements: see Appendix
Male. Unknown.
Forest leaf litter.
Known only from the type locality (Fig.
Nesticella yui Wunderlich & Song, 1995: 347, fig. 19 (♀).
Nesticella
yui
:
Nesticella
yui
:
Holotype ♀ and paratype 1♀ (
2♂ (
Males of this species can be easily distinguished from those of the congeneric members by the single, massive distal process of the paracymbium (see
See
Forest leaf litter.
China (Yunnan) (Fig.
Holotype ♂ and paratypes 1♂4♀ (
The new species is named after Dr. Zhiyuan Yao who extensively collected and studied spiders from Southeast Asia; noun (name) in genitive case.
Males of Nesticella zhiyuani sp. n. can be recognized from those of the other species belonging to the nepalensis-group by the long, sickle-like terminal apophysis (Ta) with a very sharp tip, by the reduced, blunt tegular apophysis (Tg-I) and by the almost round distal process I (Dp-I) and the elongate, sharp process II (Dp-II) (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.13. Carapace 1.15 long, 1.05 wide. Opisthosoma 1.05 long, 0.95 wide. Clypeus height 0.22. Sternum 0.65 long, 0.68 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.50. Carapace 1.14 long, 0.99 wide. Opisthosoma 1.30 long, 1.35 wide. Clypeus height 0.20. Sternum 0.66 long, 0.66 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Group features. Males of the species belonging to this group can be distinguished by having a short, flat and unbranched ventral apophysis (Va), sometimes strongly reduced, by a flat, an elongate or a squared distal process of the paracymbium (Dp), a well-developed tegular apophysis (Tg) and a hook-shaped process of the conductor (Cp). Females are easily recognized by the short fertilization and copulatory ducts (Fd and Cd) and the wide, ovoid spermathecae (S).
Composition. Nesticella phami sp. n. and N. sumatrana sp. n.
Holotype ♂ and paratypes 1♂3♀ (
The new species is named after Dr. Pham Dinh Sac, a pioneer spider researcher in Vietnam; noun (name) in genitive case.
Males of the new species can be distinguished from those of Nesticella sumatrana sp. n. (Fig.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.72. Carapace 1.00 long, 0.93 wide. Opisthosoma 0.83 long, 0.63 wide. Clypeus height 0.19. Sternum 0.60 long, 0.57 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.28. Carapace 1.10 long, 0.98 wide. Opisthosoma 1.25 long, 0.96 wide. Clypeus height 0.20. Sternum 0.68 long, 0.67 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Holotype ♂ (
The specific name is derived from the type locality; adjective.
Nesticella sumatrana sp. n. can be distinguished from N. phami sp. n. (Figs
Habitus as in Fig.
Male palp (Fig.
Male (holotype). Total length 1.81. Carapace 0.92 long, 0.85 wide. Opisthosoma 0.96 long, 0.86 wide. Clypeus height 0.17. Sternum 0.56 long, 0.57 wide. Leg measurements: see Appendix
Female. Unknown.
Cave.
Known only from the type locality (Fig.
Group features. Males belonging to this species-group can be recognized by having a paracymbium with a very wide, laminar distal process (Dp), a short ventral apophysis (Va), a protruding, crest-like terminal apophysis (Ta), a missing tegular apophysis (Tg) and a wide, flat process of the conductor (Cp). Females are distinguished by the presence of a protruding scape with a rounded apex and by the straight copulatory ducts (Cd), strongly diverging to each other and forming a V visible through the transparent tegument of the epigyne.
Composition. Nesticella kaohsiungensis sp. n. and N. quelpartensis (Paik & Namkung, 1969).
Nesticella taiwan Tso & Yoshida, 2000: 13, figs 7–11 (only ♀, mismatched with the holotype ♂).
Holotype ♂ (
The specific name is derived from the type locality; adjective.
Males can be distinguished from those of N. quelpartensis (see Paik & Namkung, in
Habitus as Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.88. Carapace 1.06 long, 0.92 wide. Opisthosoma 0.94 long, 0.68 wide. Clypeus height 0.18. Sternum 0.64 long, 0.61 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.29. Carapace 1.20 long, 1.00 wide. Opisthosoma 1.30 long, 0.93 wide. Clypeus height 0.21. Sternum 0.71 long, 0.67 wide. Leg measurements: see Appendix
Cave, forest leaf litter.
China (Taiwan) (Fig.
A female paratype of Nesticella taiwan Tso & Yoshida, 2000 was mismatched with the holotype male, synonymized here with N. odonta. A detailed morphological comparison of the samples, together with the an in-progress molecular analysis of newly collected specimens from the type locality of N. taiwan, allow the correct matching of females and males and demonstrate that they belong to a new species.
Nesticus
quelpartensis
Paik & Namkung, in
Nesticus
quelpartensis
:
Howaia
quelpartensis
:
Nesticella
quelpartensis
:
Nesticella
quelpartensis
:
Males can be separated from those of Nesticella kaohsiungensis sp. n. by the wider ventral apophysis of the paracymbium (Va) and the slimmer and sharper process of the conductor (Cp) (see
See
Cave.
Korea.
Pseudonesticus Liu & Li, 2013a: 790.
Pseudonesticus clavatus Liu & Li, 2013 from Yunnan, China.
Males belonging to Pseudonesticus can be distinguished from those of Cyclocarcina Komatsu, 1942 by the hooked or finger-like terminal apophysis (Ta), always well-developed (shorter and simpler in Cyclocarcina), the longer embolus, the flat tegular apophysis (Tg) and the less ramified paracymbium, rather than having several apophyses. It can be separated from the other Nesticini by the slimmer or hook shaped terminal apophysis (Ta) and by the usually very long embolus (E) which are thicker and shorter in the other genera. Females can be distinguished from those belonging to the other Nesticini by the shape of the fertilization and copulatory ducts, coiled before reaching the spermathecae, and by the wide or triangular scape which is not shared with any other genus.
Total length: 1.76–2.72 (male), 1.82–3.24 (female). Carapace almost round in males, ovate in females, usually uniformly yellow. Legs of the same color. Eyes generally reduced or absent. When present, AER and PER straight, MOA trapezoidal, narrower in the front. Eight well-developed eyes and a clear pattern only in P. dafangensis sp. n. Cervical groove and fovea usually indistinct. Chelicera with three promarginal teeth and multiple retromarginal tiny denticles on the fang furrow. Opisthosoma with long setae, yellowish or greyish, rarely heavily pigmented.
Male palp: tibia short, wider than long, basally narrower, with three retrolateral trichobothria and several long setae. Paracymbium well-developed with a laminar distal process, pointed or lobe-shaped, and generally lacking a ventral apophysis, which is always simple when present (e.g. P. clavatus). Some species with a long and flat dorsal apophysis. Bulb with a well-developed terminal apophysis, hooked or finger-like. Tegular apophysis flat, P. spinosus sp. n. and P. ziyunensis sp. n. with a second tegular apophysis (Tg-II). Conductor wide and laminar, with one or two processes, sometimes with a long apex (e.g. P. miao sp. n., P. ziyunensis sp. n.). Embolus usually thin and long, with the terminal part strongly coiled, reduced only in P. clavatus.
Epigyne: scape always present, wide and lobed or triangularly-shaped. Copulatory openings located under the scape. Spermathecae slightly visible through the tegument. Spermathecae small and globular separated by at least two diameters. Fertilization and copulatory ducts ventrally oriented, thin and coiled with two to three loops. Vulval pockets well developed, located near the spermathecae.
Pseudonesticus clavatus Liu & Li, 2013, P. dafangensis sp. n., P. miao sp. n., P. spinosus sp. n., P. wumengensis sp. n., and P. ziyunensis sp. n.
China (Guizhou, Yunnan).
Pseudonesticus clavatus Liu & Li, 2013a: 790, figs 1–14 (♂♀).
Males can be recognized from those of other Pseudonesticus species by the wider distal process (Dp) of the paracymbium, the big and thick process of the conductor (Cp), the wide and well-developed ventral apophysis of the paracymbium (Va) and the thick embolus which are respectively, narrower, smaller and thinner, less developed or absent and slimmer in the other species (see
See
China (Yunnan).
Holotype ♂ and paratypes 8♂14♀ (
The specific name is derived from the type locality; adjective.
The new species is closely related to Pseudonesticus spinosus sp. n. (Figs
Pseudonesticus dafangensis sp. n., holotype (male) and paratype (female). A Male habitus, dorsal view B Ditto, ventral view C Female habitus, dorsal view D Ditto, ventral view E Epigyne, ventral view F Ditto, lateral view G Vulva, ventral view H Vulva, dorsal view. Scale bars: A–D = 0.50 mm; E–H = 0.10 mm.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.76. Carapace 0.95 long, 0.81 wide. Opisthosoma 0.92 long, 0.71 wide. Clypeus height 0.20. Sternum 0.58 long, 0.55 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.07. Carapace 1.01 long, 0.80 wide. Opisthosoma 1.23 long, 0.92 wide. Clypeus length 0.18. Sternum 0.62 long, 0.48 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 18♀ (
Named after the Miao people, an ethnic minority living in Guizhou Province; noun in apposition.
The new species is closely related to P. ziyunensis sp. n. (Figs
Pseudonesticus miao sp. n, holotype (male) and paratype (female). A Male prosoma, dorsal view B Ditto, ventral view C Female habitus, dorsal view D Ditto, ventral view E Ditto, lateral view F Epigyne, ventral view G Vulva, ventral view H Vulva, dorsal view. Scale bars: A–E = 0.50 mm; F–H = 0.10 mm.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.58. Carapace 1.13 long, 1.01 wide. Opisthosoma 1.56 long, 1.05 wide. Sternum 0.65 long, 0.65 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 2.64. Carapace 1.18 long, 0.99 wide. Opisthosoma 1.60 long, 1.19 wide. Sternum 0.66 long, 0.69 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 4♀ (
The specific name is derived from the Latin word “spinosus” = spiny, thorny, and it is related to the spiked shape of the distal process of the paracymbium in the male; adjective.
The new species is closely related to P. dafangensis sp. n. (Figs
Pseudonesticus spinosus sp. n., holotype (male) and paratype (female). A Male habitus, dorsal view B Ditto, ventral view C Female habitus, dorsal view D Ditto, ventral view E Epigyne, ventral view F Ditto, lateral view G Vulva, dorsal view H Vulva, posterior view. Scale bars: A–D = 0.50 mm; E–H = 0.10 mm.
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 1.78. Carapace 0.84 long, 0.73 wide. Opisthosoma 1.03 long, 0.83 wide. Clypeus height 0.17. Sternum 0.54 long, 0.47 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 1.82. Carapace 0.74 long, 0.69 wide. Opisthosoma 1.00 long, 1.08 wide. Clypeus length 0.15. Sternum 0.50 long, 0.45 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Holotype ♀ and paratype 1♀ (
The specific name is derived from the type locality; adjective.
This species is closely related to P. dafangensis sp. n. (see Fig.
Habitus as in Fig.
Epigyne (Fig.
Female (holotype). Total length 2.13. Carapace 0.95 long, 0.77 wide. Opisthosoma 1.38 long, 1.05 wide. Clypeus height 0.20. Sternum 0.60 long, 0.55 wide. Leg measurements: see Appendix
Male. Unknown.
Cave.
Known only from the type locality (Fig.
Holotype ♂ and paratypes 2♂12♀ (
The specific name is derived from the type locality; adjective.
The new species is closely related to P. miao sp. n. (Figs
Habitus as in Fig.
Male palp (Fig.
Epigyne (Fig.
Male (holotype). Total length 2.72. Carapace 1.19 long, 1.06 wide. Opisthosoma 1.64 long, 1.00 wide. Sternum 0.66 long, 0.70 wide. Leg measurements: see Appendix
Female (one of the paratypes). Total length 3.16. Carapace 1.27 long, 1.06 wide. Opisthosoma 1.82 long, 1.24 wide. Sternum 0.70 long, 0.74 wide. Leg measurements: see Appendix
Cave.
Known only from the type locality (Fig.
Speleoticus navicellatus Liu & Li, 2013 from Guangxi, China.
The generic name is a combination of the Greek word “Speleo-” = cave and the contract name of Nesticus, the nominal genus of the family. It alludes to the troglophyllic lifestyle of these species. The gender is masculine.
Males belonging to Speleoticus gen. n. can be distinguished from those of the other Nesticini by the relatively simple, sickle-shaped paracymbium (P) with only a few short apophyses, in contrast to the other genera where it can be more complex, usually with wide and long processes. Furthermore, the elongate tibia and the triangular or rectangular protruding terminal apophysis (Ta) allow quick separation from the other Asian genera of Nesticidae. Females can be separated from those of the other Nesticini, with the exception of Cyclocarcina, by the wide, balloon-shaped vulval pockets (Vp) located above the spermathecae which are usually around or below the spermathecae in the other genera. Females of Speleoticus gen. n. can be easily separated from those of Cyclocarcina by the very short and narrow scape (Sp) which is well-developed and strongly protruding in the latter genus.
Total length: 2.84–3.15 (male), 2.97–4.36 (female). Carapace almost round in males, ovate in females, uniformly pale yellow as the legs. Six eyes in two rows, AME absent. Cervical groove and fovea indistinct. Chelicera with three promarginal teeth and multiple retromarginal tiny denticles on the fang furrow. Opisthosoma yellowish with long setae (Fig.
Male palp (Fig.
Epigyne (Fig.
Distribution records of Hamus, Nescina, and Pseudonesticus spp. in China and Southeast Asia. 1 Hamus cornutus sp. n. 2 H. kangdingensis sp. n. 3 H. luzon sp. n. 4 H. mangunensis sp. n. 5 Nescina kohi sp. n. 6 Pseudonesticus dafangensis sp. n. 7 P. miao sp. n. 8 P. spinosus sp. n. 9 P. wumengensis sp. n. 10 P. ziyunensis sp. n.
Distribution records of the Nesticella spiders belonging to the brevipes-group in China. 1 Nesticella baiseensis sp. n. 2 N. caeca sp. n. 3 N. chongqing sp. n. 4 N. dazhuangensis sp. n. 5 N. gazuida sp. n. 6 N. hongheensis sp. n. 7 N. jingpo sp. n. 8 N. lisu sp. n. 9 N. liuzhaiensis sp. n. 10 N. nandanensis sp. n. 11 N. odonta 12 N. qiaoqiensis sp. n. 13 N. qiongensis sp. n. 14 N. robusta sp. n. 15 N. sanchaheensis sp. n. 16 N. songi 17 N. xiongmao sp. n. 18 N. xixia sp. n. 19 N. yao sp. n.
Distribution records of the Nesticella spiders belonging to the nepalensis, mogera, phami and quelpartensis-groups in Asia and Madagascar. 1 Nesticella baobab sp. n. 2 N. connectens 3 N. gongshanensis sp. n. 4 N. griswoldi sp. n. 5 N. nepalensis 6 N. potala sp. n. 7 N. sulawesi sp. n. 8 N. tibetana sp. n. 9 N. vanlang sp. n. 10 N. yui Wunderlich et Song, 1995 11 N. zhiyuani sp. n. 12 N. fuliangensis sp. n. 13 Nesticella huomachongensis sp. n. 14 N. rongtangensis sp. n. 15 N. wanzaiensis sp. n. 16 N. yanbeiensis sp. n. 17 N. phami sp. n. 18 N. sumatrana sp. n. 19 N. kaohsiungensis sp. n.
Speleoticus globosus (Liu & Li, 2013), comb. n., S. libo (Chen & Zhu, 2005), comb. n., S. navicellatus (Liu & Li, 2013), comb. n., S. uenoi (Yaginuma, 1972), comb. n., and S. yaginumai (Yin, 2012), comb. n. All the species listed above are transferred from the genus Nesticus. All new combinations are supported by our molecular phylogenetic analysis.
China (Guangxi, Guizhou, Hunan), Japan (Honshu Island, Shizuoka Prefecture).
Speleoticus gen. n., together with the Pseudonesticus Liu & Li, 2013, show typical adaptations to cave life, such as the absence or reduction of the eyes (in particular the AME), long legs, lack of pigmentation, etc. The close relationship between these two genera, rather than with the genera from the West Palaearctic and North America, can be cautiously hypothesized based on the morphological comparison and preliminary molecular analysis of Nesticidae. Nevertheless, further studies are necessary to understand their correct systematic position within the family. Here we illustrate male and female of Speleoticus navicellatus, the type species of Speleoticus gen. n.
Nesticus navicellatus Liu & Li, 2013b: 542, figs 35A–B, 36A–D, 37A–D, 38A–E (♂♀).
Holotype ♂, paratype 1♀ (
1♂ (
See
See Figs
Cave.
China (Guangxi).
This study was supported by the Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences (2015CASEABRI005, Y4ZK111B01) and the National Natural Sciences Foundation of China (NSFC-31272280, 31471960, 31530067, 31672260). The manuscript benefitted greatly from comments by Yuri M. Marusik (Magadan, Russia) and Nadine Dupérré (Quito, Ecuador). Sarah Crews (Oakland, USA) kindly checked the English of the manuscript. Peter Jäger (Frankfurt, Germany) checked the proof. Lena Chan, Joseph K. H. Koh (Singapore) and colleagues of the Asian Society of Arachnology kindly helped for the collecting permission. Charles E. Griswold (San Francisco, USA) kindly allowed us to study his material collected from Madagascar. Yuri M. Marusik allowed us to check the male of Nesticella nepalensis, Andrei T. Tanasevitch (Moscow, Russia) kindly provided photos of Nesticella mollicula.
Hamus cornutus sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.23 | 1.03 | 0.76 | 1.05 | |
Patella | 0.33 | 0.31 | 0.27 | 0.30 | |
Tibia | 1.14 | 0.84 | 0.55 | 0.89 | |
Metatarsus | 0.78 | 0.68 | 0.46 | 0.63 | |
Tarsus | 0.70 | 0.65 | 0.54 | 0.65 | |
Total | 4.18 | 3.51 | 2.58 | 3.52 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.25 | 1.06 | 0.79 | 1.19 | 0.35 |
Patella | 0.36 | 0.36 | 0.31 | 0.35 | 0.15 |
Tibia | 1.08 | 0.87 | 0.55 | 1.00 | 0.19 |
Metatarsus | 0.79 | 0.68 | 0.47 | 0.66 | ‒ |
Tarsus | 0.69 | 0.66 | 0.52 | 0.64 | 0.41 |
Total | 4.17 | 3.63 | 2.64 | 3.84 | 1.10 |
Hamus kangdingensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.12 | 0.94 | 0.70 | 1.05 | |
Patella | 0.30 | 0.28 | 0.24 | 0.28 | |
Tibia | 0.92 | 0.72 | 0.49 | 0.84 | |
Metatarsus | 0.63 | 0.53 | 0.40 | 0.53 | |
Tarsus | 0.64 | 0.57 | 0.47 | 0.59 | |
Total | 3.61 | 3.04 | 2.30 | 3.29 | |
Hamus luzon sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.21 | 1.02 | 0.80 | 1.22 | 0.39 |
Patella | 0.39 | 0.36 | 0.32 | 0.35 | 0.17 |
Tibia | 0.99 | 0.80 | 0.58 | 1.01 | 0.21 |
Metatarsus | 0.71 | 0.61 | 0.47 | 0.65 | ‒ |
Tarsus | 0.62 | 0.57 | 0.52 | 0.61 | 0.42 |
Total | 3.92 | 3.36 | 2.69 | 3.84 | 1.19 |
Nescina kohi sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 0.67 | 0.57 | 0.50 | 0.61 | |
Patella | 0.23 | 0.20 | 0.19 | 0.22 | |
Tibia | 0.54 | 0.46 | 0.32 | 0.47 | |
Metatarsus | 0.37 | 0.32 | 0.29 | 0.35 | |
Tarsus | 0.31 | 0.26 | 0.25 | 0.28 | |
Total | 2.12 | 1.81 | 1.55 | 1.93 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 0.68 | 0.62 | 0.53 | 0.64 | 0.33 |
Patella | 0.22 | 0.22 | 0.19 | 0.21 | 0.18 |
Tibia | 0.53 | 0.46 | 0.33 | 0.49 | 0.23 |
Metatarsus | 0.36 | 0.33 | 0.30 | 0.35 | ‒ |
Tarsus | 0.30 | 0.29 | 0.26 | 0.27 | 0.40 |
Total | 2.09 | 1.92 | 1.61 | 1.96 | 1.14 |
Nesticella baiseensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.90 | 2.37 | 1.92 | 2.60 | |
Patella | 0.65 | 0.60 | 0.52 | 0.61 | |
Tibia | 2.71 | 1.98 | 1.40 | 2.08 | |
Metatarsus | 2.68 | 2.00 | 1.12 | 1.92 | |
Tarsus | 1.15 | 0.94 | 0.76 | 0.88 | |
Total | 10.09 | 7.89 | 5.72 | 8.09 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.55 | 2.00 | 1.60 | 2.21 | 0.69 |
Patella | 0.64 | 0.58 | 0.52 | 0.61 | 0.24 |
Tibia | 2.32 | 1.60 | 1.08 | 1.76 | 0.35 |
Metatarsus | 2.08 | 1.52 | 1.14 | 1.48 | ‒ |
Tarsus | 1.02 | 0.88 | 0.71 | 0.82 | 0.81 |
Total | 8.61 | 6.58 | 5.05 | 6.88 | 2.09 |
Nesticella caeca sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.43 | 1.99 | 1.60 | 2.15 | |
Patella | 0.53 | 0.50 | 0.41 | 0.45 | |
Tibia | 2.30 | 1.68 | 1.18 | 1.78 | |
Metatarsus | 2.32 | 1.67 | 1.20 | 1.63 | |
Tarsus | 1.04 | 0.85 | 0.73 | 0.83 | |
Total | 8.62 | 6.69 | 5.12 | 6.84 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.45 | 1.98 | 1.58 | 2.19 | 0.65 |
Patella | 0.55 | 0.48 | 0.41 | 0.50 | 0.23 |
Tibia | 2.23 | 1.63 | 1.06 | 1.72 | 0.32 |
Metatarsus | 2.10 | 1.50 | 1.10 | 1.44 | ‒ |
Tarsus | 1.05 | 0.90 | 0.72 | 0.84 | 0.75 |
Total | 8.38 | 6.49 | 4.87 | 6.69 | 1.95 |
Nesticella chongqing sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.98 | ‒ | 1.14 | 1.75 | 0.55 |
Patella | 0.53 | ‒ | 0.40 | 0.50 | 0.23 |
Tibia | 1.75 | ‒ | 0.78 | 1.35 | 0.29 |
Metatarsus | 1.63 | ‒ | 0.85 | 1.15 | ‒ |
Tarsus | 0.78 | ‒ | 0.55 | 0.63 | 0.64 |
Total | 6.67 | ‒ | 3.72 | 5.38 | 1.71 |
Nesticella dazhuangensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.08 | 1.70 | 1.40 | 1.90 | |
Patella | 0.58 | 0.53 | 0.44 | 0.53 | |
Tibia | 1.83 | 1.35 | 0.96 | 1.53 | |
Metatarsus | 1.93 | 1.36 | 1.06 | 1.48 | |
Tarsus | 0.80 | 0.70 | 0.62 | 0.73 | |
Total | 7.22 | 5.64 | 4.48 | 6.17 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.72 | 1.38 | 1.05 | 1.58 | 0.52 |
Patella | 0.51 | 0.43 | 0.40 | 0.50 | 0.21 |
Tibia | 1.55 | 1.13 | 0.75 | 1.25 | 0.27 |
Metatarsus | 1.44 | 1.04 | 0.80 | 1.10 | ‒ |
Tarsus | 0.72 | 0.63 | 0.55 | 0.60 | 0.60 |
Total | 5.94 | 4.61 | 3.55 | 5.03 | 1.60 |
Nesticella gazuida sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.47 | 2.00 | 1.58 | 2.16 | 0.65 |
Patella | 0.56 | 0.50 | 0.44 | 0.52 | 0.23 |
Tibia | 2.25 | 1.66 | 1.06 | 1.73 | 0.34 |
Metatarsus | 2.13 | 1.56 | 1.13 | 1.53 | ‒ |
Tarsus | 0.88 | 0.78 | 0.69 | 0.75 | 0.73 |
Total | 8.29 | 6.50 | 4.90 | 6.69 | 1.95 |
Nesticella hongheensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.88 | 1.44 | 1.15 | 1.63 | |
Patella | 0.50 | 0.44 | 0.38 | 0.41 | |
Tibia | 1.72 | 1.16 | 0.78 | 1.31 | |
Metatarsus | 1.78 | 1.19 | 0.94 | 1.25 | |
Tarsus | 0.77 | 0.64 | 0.52 | 0.62 | |
Total | 6.65 | 4.87 | 3.77 | 5.22 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.82 | 1.38 | 1.11 | 1.62 | 0.54 |
Patella | 0.51 | 0.48 | 0.38 | 0.46 | 0.22 |
Tibia | 1.54 | 1.02 | 0.70 | 1.22 | 0.28 |
Metatarsus | 1.45 | 1.03 | 0.79 | 1.04 | ‒ |
Tarsus | 0.73 | 0.61 | 0.51 | 0.58 | 0.60 |
Total | 6.05 | 4.52 | 3.49 | 4.92 | 1.64 |
Nesticella jingpo sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.48 | 1.18 | 0.93 | 1.35 | 0.44 |
Patella | 0.43 | 0.40 | 0.38 | 0.41 | 0.18 |
Tibia | 1.33 | 0.90 | 0.63 | 1.08 | 0.22 |
Metatarsus | 1.08 | 0.88 | 0.69 | 0.93 | ‒ |
Tarsus | 0.70 | 0.56 | 0.49 | 0.55 | 0.53 |
Total | 5.02 | 3.92 | 3.12 | 4.32 | 1.37 |
Nesticella lisu sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.85 | 1.45 | 1.15 | 1.66 | |
Patella | 0.49 | 0.46 | 0.38 | 0.43 | |
Tibia | 1.69 | 1.14 | 0.80 | 1.35 | |
Metatarsus | 1.68 | 1.13 | 0.83 | 1.25 | |
Tarsus | 0.75 | 0.65 | 0.55 | 0.65 | |
Total | 6.46 | 4.83 | 3.71 | 5.34 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.75 | 1.41 | 1.15 | 1.63 | 0.54 |
Patella | 0.49 | 0.47 | 0.38 | 0.48 | 0.21 |
Tibia | 1.54 | 1.06 | 0.75 | 1.29 | 0.27 |
Metatarsus | 1.40 | 1.00 | 0.81 | 1.05 | ‒ |
Tarsus | 0.71 | 0.63 | 0.56 | 0.65 | 0.59 |
Total | 5.89 | 4.57 | 3.65 | 5.10 | 1.61 |
Nesticella liuzhaiensis sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.53 | 1.18 | 0.93 | 1.40 | 0.40 |
Patella | 0.40 | 0.37 | 0.33 | 0.38 | 0.19 |
Tibia | 1.33 | 0.89 | 0.62 | 1.35 | 0.21 |
Metatarsus | 1.18 | 0.81 | 0.63 | 0.88 | ‒ |
Tarsus | 0.64 | 0.55 | 0.49 | 0.58 | 0.51 |
Total | 5.08 | 3.80 | 3.00 | 4.59 | 1.31 |
Nesticella nandanensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.30 | 1.81 | 1.53 | 2.06 | |
Patella | 0.59 | 0.53 | 0.45 | 0.53 | |
Tibia | 2.13 | 1.50 | 1.10 | 1.69 | |
Metatarsus | 2.19 | 1.56 | 1.21 | 1.59 | |
Tarsus | 0.94 | 0.78 | 0.65 | 0.75 | |
Total | 8.15 | 6.18 | 4.94 | 6.62 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.44 | 1.91 | 1.52 | 2.15 | 0.71 |
Patella | 0.66 | 0.60 | 0.49 | 0.61 | 0.25 |
Tibia | 2.23 | 1.53 | 1.03 | 1.73 | 0.36 |
Metatarsus | 2.08 | 1.50 | 1.09 | 1.49 | ‒ |
Tarsus | 0.91 | 0.78 | 0.65 | 0.74 | 0.80 |
Total | 8.32 | 6.32 | 4.78 | 6.72 | 2.12 |
Nesticella qiaoqiensis sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | ‒ | 1.53 | 1.16 | 1.72 | 0.55 |
Patella | ‒ | 0.47 | 0.41 | 0.49 | 0.21 |
Tibia | ‒ | 1.19 | 0.75 | 1.28 | 0.27 |
Metatarsus | ‒ | 1.14 | 0.84 | 1.13 | ‒ |
Tarsus | ‒ | 0.69 | 0.59 | 0.63 | 0.63 |
Total | ‒ | 5.02 | 3.75 | 5.25 | 1.66 |
Nesticella qiongensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.93 | 1.42 | 1.13 | 1.62 | |
Patella | 0.45 | 0.40 | 0.33 | 0.42 | |
Tibia | 1.83 | 1.16 | 0.78 | 1.36 | |
Metatarsus | 1.85 | 1.14 | 0.87 | 1.28 | |
Tarsus | 0.74 | 0.60 | 0.52 | 0.62 | |
Total | 6.80 | 4.72 | 3.63 | 5.30 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.60 | 1.21 | 0.95 | 1.58 | 0.45 |
Patella | 0.46 | 0.41 | 0.35 | 0.45 | 0.19 |
Tibia | 1.36 | 0.89 | 0.59 | 1.34 | 0.25 |
Metatarsus | 1.24 | 0.86 | 0.67 | 1.22 | ‒ |
Tarsus | 0.64 | 0.56 | 0.48 | 0.63 | 0.53 |
Total | 5.30 | 3.93 | 3.04 | 5.22 | 1.42 |
Nesticella robusta sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.03 | 1.47 | 1.15 | 1.66 | |
Patella | 0.46 | 0.42 | 0.34 | 0.43 | |
Tibia | 1.87 | 1.18 | 0.80 | 1.39 | |
Metatarsus | 1.84 | 1.15 | 0.91 | 1.30 | |
Tarsus | 0.76 | 0.61 | 0.54 | 0.63 | |
Tatal | 6.96 | 4.83 | 3.74 | 5.41 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.82 | 1.38 | 1.08 | 1.67 | 0.46 |
Patella | 0.47 | 0.44 | 0.33 | 0.45 | 0.19 |
Tibia | 1.38 | 1.12 | 0.78 | 1.34 | 0.26 |
Metatarsus | 1.25 | 1.08 | 0.73 | 1.23 | ‒ |
Tarsus | 0.68 | 0.60 | 0.50 | 0.63 | 0.55 |
Tatal | 5.60 | 4.62 | 3.42 | 5.32 | 1.46 |
Nesticella sanchaheensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.84 | 2.34 | ‒ | ‒ | |
Patella | 0.59 | 0.53 | ‒ | ‒ | |
Tibia | 2.56 | 1.69 | ‒ | ‒ | |
Metatarsus | 2.50 | 1.88 | ‒ | ‒ | |
Tarsus | 1.09 | 0.88 | ‒ | ‒ | |
Total | 9.58 | 7.32 | ‒ | ‒ | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 3.16 | 2.75 | 2.04 | 2.64 | 0.78 |
Patella | 0.64 | 0.61 | 0.48 | 0.60 | 0.26 |
Tibia | 2.90 | 2.15 | 1.39 | 2.20 | 0.40 |
Metatarsus | 2.72 | 1.90 | 1.48 | 2.06 | ‒ |
Tarsus | 1.20 | 0.93 | 0.79 | 1.04 | 0.89 |
Total | 10.62 | 8.34 | 6.18 | 8.54 | 2.33 |
Nesticella xiongmao sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.19 | 1.63 | 1.28 | 1.84 | |
Patella | 0.48 | 0.47 | 0.41 | 0.48 | |
Tibia | 2.05 | 1.00 | 0.91 | 1.48 | |
Metatarsus | 1.97 | 1.27 | 0.97 | 1.36 | |
Tarsus | 0.80 | 0.66 | 0.53 | 0.63 | |
Total | 7.49 | 5.03 | 4.10 | 5.79 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.83 | 1.38 | 1.12 | 1.63 | 0.52 |
Patella | 0.50 | 0.46 | 0.38 | 0.48 | 0.22 |
Tibia | 1.65 | 1.08 | 0.73 | 1.26 | 0.26 |
Metatarsus | 1.48 | 1.00 | 0.78 | 1.05 | ‒ |
Tarsus | 0.73 | 0.63 | 0.53 | 0.64 | 0.58 |
Total | 6.19 | 4.55 | 3.54 | 5.06 | 1.58 |
Nesticella xixia sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 3.00 | 2.32 | 1.85 | 2.55 | |
Patella | 0.68 | 0.64 | 0.53 | 0.59 | |
Tibia | 2.76 | 1.91 | 1.33 | 2.03 | |
Metatarsus | 2.68 | 1.88 | 1.45 | 1.91 | |
Tarsus | 1.04 | 0.91 | 0.73 | 0.88 | |
Total | 10.16 | 7.66 | 5.89 | 7.96 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.88 | 2.20 | 1.80 | 2.50 | 0.78 |
Patella | 0.64 | 0.60 | 0.52 | 0.63 | 0.22 |
Tibia | 2.68 | 1.81 | 1.18 | 1.94 | 0.41 |
Metatarsus | 2.36 | 1.68 | 1.31 | 1.63 | ‒ |
Tarsus | 1.00 | 0.85 | 0.69 | 0.78 | 0.88 |
Total | 9.56 | 7.14 | 5.50 | 7.48 | 2.29 |
Nesticella yao sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.35 | 1.81 | 1.50 | 2.03 | 0.66 |
Patella | 0.56 | 0.53 | 0.48 | 0.53 | 0.23 |
Tibia | 2.09 | 1.45 | 0.98 | 1.63 | 0.32 |
Metatarsus | 1.94 | 1.38 | 1.08 | 1.34 | ‒ |
Tarsus | 0.91 | 0.78 | 0.65 | 0.70 | 0.76 |
Total | 7.85 | 5.95 | 4.69 | 6.23 | 1.97 |
Nesticella fuliangensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.66 | 2.13 | 1.75 | 2.47 | |
Patella | 0.66 | 0.63 | 0.53 | 0.63 | |
Tibia | 2.59 | 1.88 | 1.38 | 2.22 | |
Metatarsus | 2.61 | 1.91 | 1.47 | 2.16 | |
Tarsus | 1.00 | 0.88 | 0.75 | 0.91 | |
Total | 9.52 | 7.43 | 5.88 | 8.39 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.63 | 2.06 | 1.72 | 2.38 | 0.80 |
Patella | 0.69 | 0.63 | 0.56 | 0.63 | 0.30 |
Tibia | 2.44 | 1.75 | 1.25 | 2.07 | 0.50 |
Metatarsus | 2.34 | 1.69 | 1.30 | 1.75 | ‒ |
Tarsus | 1.00 | 0.88 | 0.78 | 0.89 | 0.90 |
Total | 9.10 | 7.01 | 5.61 | 7.72 | 2.50 |
Nesticella huomachongensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.34 | 1.84 | 1.59 | 2.19 | |
Patella | 0.63 | 0.56 | 0.47 | 0.59 | |
Tibia | 2.28 | 1.66 | 1.22 | 1.95 | |
Metatarsus | 2.33 | 1.69 | 1.31 | 1.91 | |
Tarsus | 1.04 | 0.88 | 0.72 | 0.89 | |
Total | 8.62 | 6.63 | 5.31 | 7.53 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.32 | 1.84 | 1.56 | 2.16 | 0.70 |
Patella | 0.64 | 0.60 | 0.51 | 0.59 | 0.29 |
Tibia | 2.12 | 1.53 | 1.09 | 1.63 | 0.35 |
Metatarsus | 2.06 | 1.52 | 1.16 | 1.56 | ‒ |
Tarsus | 1.00 | 0.88 | 0.75 | 0.83 | 0.87 |
Total | 8.14 | 6.37 | 5.07 | 6.77 | 2.21 |
Nesticella rongtangensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.09 | 1.59 | 1.34 | 1.88 | |
Patella | 0.47 | 0.41 | 0.35 | 0.42 | |
Tibia | 2.03 | 1.44 | 1.03 | 1.69 | |
Metatarsus | 2.00 | 1.38 | 1.06 | 1.53 | |
Tarsus | 0.84 | 0.66 | 0.56 | 0.71 | |
Total | 7.43 | 5.48 | 4.34 | 6.23 | |
Nesticella wanzaiensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.28 | 1.72 | 1.50 | 2.00 | |
Patella | 0.61 | 0.53 | 0.45 | 0.50 | |
Tibia | 2.23 | 1.59 | 1.14 | 1.81 | |
Metatarsus | 2.31 | 1.56 | 1.22 | 1.69 | |
Tarsus | 1.06 | 0.83 | 0.72 | 0.84 | |
Total | 8.49 | 6.23 | 5.03 | 6.84 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.36 | 1.77 | 1.56 | 2.13 | |
Patella | 0.65 | 0.62 | 0.49 | 0.60 | |
Tibia | 2.31 | 1.65 | 1.18 | 1.92 | |
Metatarsus | 2.33 | 1.60 | 1.25 | 1.73 | |
Tarsus | 1.12 | 0.93 | 0.80 | 0.88 | |
Total | 8.77 | 6.57 | 5.28 | 7.26 | |
Nesticella yanbeiensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.22 | 1.83 | 1.53 | 2.00 | |
Patella | 0.66 | 0.60 | 0.48 | 0.53 | |
Tibia | 2.05 | 1.53 | 1.15 | 1.78 | |
Metatarsus | 2.06 | 1.55 | 1.23 | 1.66 | |
Tarsus | 1.03 | 0.91 | 0.73 | 0.91 | |
Total | 8.02 | 6.42 | 5.12 | 6.88 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.31 | 1.91 | 1.56 | 2.15 | 0.72 |
Patella | 0.66 | 0.63 | 0.50 | 0.58 | 0.25 |
Tibia | 2.13 | 1.53 | 1.10 | 1.80 | 0.38 |
Metatarsus | 2.00 | 1.47 | 1.11 | 1.58 | ‒ |
Tarsus | 0.97 | 0.88 | 0.70 | 0.85 | 0.82 |
Total | 8.07 | 6.42 | 4.97 | 6.96 | 2.17 |
Nesticella baobab sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.55 | 1.23 | 0.96 | 1.43 | |
Patella | 0.44 | 0.40 | 0.33 | 0.39 | |
Tibia | 1.48 | 1.01 | 0.68 | 1.19 | |
Metatarsus | 1.40 | 0.93 | 0.74 | 1.38 | |
Tarsus | 0.63 | 0.53 | 0.45 | 0.55 | |
Total | 5.50 | 4.10 | 3.16 | 4.94 | |
Nesticella connectens Wunderlich, 1995 | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.98 | 1.64 | 1.42 | 1.76 | |
Patella | 0.55 | 0.51 | 0.49 | 0.52 | |
Tibia | 1.83 | 1.41 | 0.96 | 1.48 | |
Metatarsus | 1.75 | 1.35 | 0.92 | 1.43 | |
Tarsus | 0.98 | 0.84 | 0.58 | 0.83 | |
Total | 7.09 | 5.75 | 4.37 | 6.02 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.94 | 1.58 | 1.28 | 1.80 | 0.63 |
Patella | 0.54 | 0.49 | 0.46 | 0.51 | 0.24 |
Tibia | 1.77 | 1.24 | 0.87 | 1.42 | 0.34 |
Metatarsus | 1.62 | 1.22 | 0.98 | 1.37 | ‒ |
Tarsus | 0.77 | 0.66 | 0.57 | 0.65 | 0.73 |
Total | 6.64 | 5.19 | 4.16 | 5.75 | 1.94 |
Nesticella gongshanensis sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | ‒ | 1.44 | 1.14 | 1.64 | 0.54 |
Patella | ‒ | 0.45 | 0.31 | 0.44 | 0.22 |
Tibia | ‒ | 1.12 | 0.76 | 1.27 | 0.30 |
Metatarsus | ‒ | 1.07 | 0.81 | 1.14 | ‒ |
Tarsus | ‒ | 0.63 | 0.52 | 0.62 | 0.59 |
Total | ‒ | 4.71 | 3.54 | 5.11 | 1.65 |
Nesticella griswoldi sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.60 | 1.25 | 0.98 | 1.41 | |
Patella | 0.48 | 0.43 | 0.35 | 0.40 | |
Tibia | 1.50 | 1.05 | 0.67 | 1.17 | |
Metatarsus | 1.50 | 1.00 | 0.66 | 0.90 | |
Tarsus | 0.81 | 0.56 | 0.40 | 0.49 | |
Total | 5.89 | 4.29 | 3.06 | 4.37 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.49 | 1.14 | 0.86 | 1.34 | 0.42 |
Patella | 0.44 | 0.40 | 0.35 | 0.42 | ‒ |
Tibia | 1.29 | 0.86 | 0.56 | 1.02 | ‒ |
Metatarsus | 1.19 | 0.84 | 0.62 | 0.96 | ‒ |
Tarsus | 0.60 | 0.50 | 0.45 | 0.52 | ‒ |
Total | 5.01 | 3.74 | 2.84 | 4.26 | ‒ |
Nesticella potala sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.56 | 2.09 | 1.65 | ‒ | 0.73 |
Patella | 0.59 | 0.57 | 0.51 | ‒ | 0.26 |
Tibia | 2.34 | 1.72 | 1.13 | ‒ | 0.36 |
Metatarsus | 2.33 | 1.42 | 1.26 | ‒ | ‒ |
Tarsus | 0.91 | 1.10 | 0.70 | ‒ | 0.89 |
Total | 8.73 | 6.90 | 5.25 | ‒ | 2.24 |
Nesticella sulawesi sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.76 | 1.40 | ‒ | 1.64 | 0.54 |
Patella | 0.52 | 0.48 | ‒ | 0.48 | 0.21 |
Tibia | 1.58 | 1.16 | ‒ | 1.33 | 0.29 |
Metatarsus | 1.47 | 1.10 | ‒ | 1.22 | ‒ |
Tarsus | 0.76 | 0.66 | ‒ | 0.68 | 0.64 |
Total | 6.09 | 4.80 | 0.00 | 5.35 | 1.68 |
Nesticella tibetana sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.64 | 1.36 | 1.09 | 1.59 | |
Patella | 0.43 | 0.41 | 0.34 | 0.46 | |
Tibia | 1.28 | 1.14 | 0.75 | 1.22 | |
Metatarsus | 1.32 | 1.36 | 0.84 | 1.10 | |
Tarsus | 0.59 | 0.58 | 0.48 | 0.62 | |
Total | 5.26 | 4.85 | 3.50 | 4.99 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.58 | 1.24 | 1.01 | 1.52 | 0.49 |
Patella | 0.49 | 0.46 | 0.38 | 0.44 | 0.20 |
Tibia | 1.37 | 0.92 | 0.64 | 1.10 | 0.25 |
Metatarsus | 1.28 | 0.93 | 0.71 | 1.06 | ‒ |
Tarsus | 0.64 | 0.56 | 0.50 | 0.62 | 0.55 |
Total | 5.36 | 4.11 | 3.24 | 4.74 | 1.49 |
Nesticella vanlang sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.68 | 1.29 | 1.00 | ‒ | 0.48 |
Patella | 0.50 | 0.43 | 0.38 | ‒ | 0.19 |
Tibia | 1.50 | 1.04 | 0.63 | ‒ | 0.25 |
Metatarsus | 1.45 | 1.00 | 0.73 | ‒ | ‒ |
Tarsus | 0.73 | 0.59 | 0.50 | ‒ | 0.57 |
Total | 5.86 | 4.35 | 3.24 | ‒ | 1.49 |
Nesticella zhiyuani sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.76 | 1.43 | 1.11 | 1.64 | |
Patella | 0.50 | 0.46 | 0.38 | 0.44 | |
Tibia | 1.70 | 1.22 | 0.81 | 1.41 | |
Metatarsus | 1.66 | 1.21 | 0.89 | 1.40 | |
Tarsus | 0.73 | 0.62 | 0.48 | 0.63 | |
Total | 6.35 | 4.94 | 3.67 | 5.52 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.39 | 1.19 | 0.97 | 1.14 | 0.48 |
Patella | 0.46 | 0.41 | 0.37 | 0.42 | 0.19 |
Tibia | 1.36 | 0.97 | 0.65 | 1.15 | 0.27 |
Metatarsus | 1.26 | 0.90 | 0.70 | 1.04 | ‒ |
Tarsus | 0.67 | 0.56 | 0.48 | 0.56 | 0.55 |
Total | 5.14 | 4.03 | 3.17 | 4.31 | 1.49 |
Nesticella phami sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.50 | 1.24 | 1.00 | 1.42 | |
Patella | 0.41 | 0.40 | 0.34 | 0.40 | |
Tibia | 1.30 | 0.94 | 0.68 | 1.12 | |
Metatarsus | 1.44 | 1.03 | 0.77 | 1.18 | |
Tarsus | 0.62 | 0.57 | 0.48 | 0.56 | |
Total | 5.27 | 4.18 | 3.27 | 4.68 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.69 | 1.40 | 1.11 | 1.60 | 0.52 |
Patella | 0.48 | 0.46 | 0.38 | 0.46 | 0.22 |
Tibia | 1.44 | 1.05 | 0.73 | 1.24 | 0.27 |
Metatarsus | 1.42 | 1.11 | 0.79 | 1.19 | ‒ |
Tarsus | 0.70 | 0.62 | 0.54 | 0.63 | 0.61 |
Total | 5.73 | 4.64 | 3.55 | 5.12 | 1.62 |
Nesticella sumatrana sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.43 | 1.20 | 1.03 | 1.28 | |
Patella | 0.38 | 0.37 | 0.30 | 0.37 | |
Tibia | 1.13 | 0.88 | 0.72 | 1.05 | |
Metatarsus | 1.05 | 0.85 | 0.75 | 1.00 | |
Tarsus | 0.55 | 0.51 | 0.49 | 0.53 | |
Total | 4.54 | 3.81 | 3.29 | 4.23 | |
Nesticella kaohsiungensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.81 | 1.52 | 1.16 | 1.66 | |
Patella | 0.47 | 0.44 | 0.36 | 0.44 | |
Tibia | 1.72 | 1.19 | 0.83 | 1.34 | |
Metatarsus | 1.75 | 1.22 | 0.90 | 1.30 | |
Tarsus | 0.75 | 0.66 | 0.53 | 0.64 | |
Total | 6.50 | 5.03 | 3.78 | 5.38 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.78 | 1.34 | 1.16 | 1.69 | 0.54 |
Patella | 0.47 | 0.44 | 0.39 | 0.46 | 0.22 |
Tibia | 1.63 | 1.09 | 0.73 | 1.32 | 0.28 |
Metatarsus | 1.50 | 1.08 | 0.81 | 1.13 | ‒ |
Tarsus | 0.73 | 0.61 | 0.51 | 0.59 | 0.63 |
Total | 6.11 | 4.56 | 3.60 | 5.19 | 1.67 |
Pseudonesticus dafangensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.44 | 1.19 | 0.98 | 1.43 | |
Patella | 0.40 | 0.38 | 0.31 | 0.37 | |
Tibia | 1.41 | 0.98 | 0.70 | 1.25 | |
Metatarsus | 1.28 | 1.05 | 0.74 | 1.06 | |
Tarsus | 0.76 | 0.65 | 0.52 | 0.65 | |
Total | 5.29 | 4.25 | 3.25 | 4.76 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.40 | 1.14 | 0.97 | 1.39 | 0.44 |
Patella | 0.43 | 0.38 | 0.35 | 0.41 | 0.18 |
Tibia | 1.31 | 0.93 | 0.66 | 1.22 | 0.28 |
Metatarsus | 1.13 | 0.84 | 0.72 | 0.94 | ‒ |
Tarsus | 0.75 | 0.63 | 0.54 | 0.62 | 0.56 |
Total | 5.02 | 3.92 | 3.24 | 4.58 | 1.46 |
Pseudonesticus miao sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 3.50 | 2.90 | 2.23 | 3.00 | |
Patella | 0.49 | 0.47 | 0.40 | 0.47 | |
Tibia | 3.70 | 2.72 | 1.73 | 2.57 | |
Metatarsus | 3.62 | 2.70 | 1.78 | 2.35 | |
Tarsus | 1.35 | 1.15 | 0.83 | 0.98 | |
Total | 12.66 | 9.94 | 6.97 | 9.37 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 3.35 | 2.64 | 2.23 | 2.81 | 0.76 |
Patella | 0.52 | 0.50 | 0.41 | 0.47 | 0.22 |
Tibia | 3.24 | 2.36 | 1.45 | 2.23 | 0.41 |
Metatarsus | 3.00 | 2.23 | 1.54 | 1.97 | ‒ |
Tarsus | 1.22 | 1.03 | 0.75 | 0.91 | 0.86 |
Total | 11.33 | 8.76 | 6.38 | 8.39 | 2.25 |
Pseudonesticus spinosus sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 1.88 | 1.53 | 1.23 | 1.78 | |
Patella | 0.36 | 0.34 | 0.31 | 0.34 | |
Tibia | 1.87 | 1.38 | 0.91 | 1.56 | |
Metatarsus | 1.74 | 1.31 | 1.01 | 1.34 | |
Tarsus | 0.90 | 0.81 | 0.66 | 0.75 | |
Total | 6.75 | 5.37 | 4.12 | 5.77 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.72 | 1.33 | 1.08 | 1.63 | 0.43 |
Patella | 0.39 | 0.35 | 0.31 | 0.36 | 0.17 |
Tibia | 1.60 | 1.13 | 0.79 | 1.33 | 0.24 |
Metatarsus | 1.46 | 1.09 | 0.83 | 1.10 | ‒ |
Tarsus | 0.88 | 0.75 | 0.60 | 0.70 | 0.5 |
Total | 6.05 | 4.65 | 3.61 | 5.12 | 1.34 |
Pseudonesticus wumengensis sp. n. | |||||
Female | |||||
I | II | III | IV | Palp | |
Femur | 1.65 | 1.43 | 1.14 | 1.66 | 0.47 |
Patella | 0.39 | 0.38 | 0.36 | 0.38 | 0.16 |
Tibia | 1.38 | 1.23 | 0.82 | 1.41 | 0.27 |
Metatarsus | 1.08 | 1.08 | 0.80 | 1.12 | ‒ |
Tarsus | 0.73 | 0.72 | 0.63 | 0.70 | 0.52 |
Total | 5.23 | 4.84 | 3.75 | 5.27 | 1.42 |
Pseudonesticus ziyunensis sp. n. | |||||
Male | |||||
I | II | III | IV | ||
Femur | 2.75 | 2.35 | 2.00 | 2.55 | |
Patella | 0.53 | 0.48 | 0.43 | 0.48 | |
Tibia | 2.69 | 2.13 | 1.50 | 2.29 | |
Metatarsus | 2.63 | 2.11 | 1.65 | 2.03 | |
Tarsus | 1.20 | 1.00 | 0.76 | 0.91 | |
Total | 9.80 | 8.07 | 6.34 | 8.26 | |
Female | |||||
I | II | III | IV | Palp | |
Femur | 2.92 | 2.38 | 1.99 | 2.66 | 0.83 |
Patella | 0.57 | 0.56 | 0.48 | 0.52 | 0.25 |
Tibia | 2.84 | 2.16 | 1.48 | 2.28 | 0.46 |
Metatarsus | 2.68 | 2.05 | 1.67 | 2.00 | ‒ |
Tarsus | 1.26 | 1.04 | 0.73 | 0.95 | 0.89 |
Total | 10.27 | 8.19 | 6.35 | 8.41 | 2.43 |
Uncorrected genetic pairwise distance of the COI partial sequence between the species discussed in this text.
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | 24 | 25 | 26 | 27 | 28 | 29 | 30 | 31 | 32 | 33 | 34 | 35 | 36 | 37 | 38 | 39 | 40 | 41 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | Hamus bowoensis | |||||||||||||||||||||||||||||||||||||||||
2 | Hamus cornutus sp. n. | 0.215 | ||||||||||||||||||||||||||||||||||||||||
3 | Hamus luzon sp. n. | 0.163 | 0.236 | |||||||||||||||||||||||||||||||||||||||
4 | Nescina kohi sp. n. | 0.248 | 0.234 | 0.234 | ||||||||||||||||||||||||||||||||||||||
5 | Nescina minuta | 0.232 | 0.234 | 0.228 | 0.097 | |||||||||||||||||||||||||||||||||||||
6 | Nesticella baiseensis sp. n. | 0.190 | 0.206 | 0.218 | 0.218 | 0.204 |