A survey of the spider family Nesticidae (Arachnida, Araneae) in Asia and Madagascar, with the description of forty-three new species

Abstract 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., Speleoticus libo (Chen & Zhu, 2005), comb. n., Speleoticus navicellatus (Liu & Li, 2015), comb. n. and Speleoticus uenoi (Yaginuma, 1972), comb. n. The new species described in this paper belong to four genera and are: Hamus cornutus sp. n. (♂♀), Hamus kangdingensis sp. n. (♂), Hamus luzon sp. n. (♀), Hamus mangunensis sp. n. (♂), Nescina kohi sp. n. (♂♀), Nesticella baiseensis sp. n. (♂♀), Nesticella baobab sp. n. (♂), Nesticella caeca sp. n. (♂♀), Nesticella chongqing sp. n. (♀), Nesticella dazhuangensis sp. n. (♂♀), Nesticella fuliangensis sp. n. (♂♀), Nesticella gazuida sp. n. (♀), Nesticella gongshanensis sp. n. (♀), Nesticella griswoldi sp. n. (♂♀), Nesticella hongheensis sp. n. (♂♀), Nesticella huomachongensis sp. n. (♂♀), Nesticella jingpo sp. n. (♀), Nesticella kaohsiungensis sp. n. (♂♀), Nesticella lisu sp. n. (♂♀), Nesticella liuzhaiensis sp. n. (♀), Nesticella nandanensis sp. n. (♂♀), Nesticella phami sp. n. (♂♀), Nesticella potala sp. n. (♀), Nesticella qiaoqiensis sp. n. (♀), Nesticella qiongensis sp. n. (♂♀), Nesticella robusta sp. n. (♂♀), Nesticella rongtangensis sp. n. (♂), Nesticella sanchaheensis sp. n. (♂♀), Nesticella sulawesi sp. n. (♀), Nesticella sumatrana sp. n. (♂), Nesticella tibetana sp. n. (♂♀), Nesticella vanlang sp. n. (♀), Nesticella wanzaiensis sp. n. (♂♀), Nesticella xiongmao sp. n. (♂♀), Nesticella xixia sp. n. (♂♀), Nesticella yanbeiensis sp. n. (♂♀), Nesticella yao sp. n. (♀), Nesticella zhiyuani sp. n. (♂♀), Pseudonesticus dafangensis sp. n. (♂♀), Pseudonesticus miao sp. n. (♂♀), Pseudonesticus spinosus sp. n. (♂♀), Pseudonesticus wumengensis sp. n. (♀), Pseudonesticus ziyunensis sp. n. (♂♀). Nesticella inthanoni (Lehtinen & Saaristo, 1980), syn. n. is synonymised with Nesticella mollicula (Thorell, 1898); Nesticella taiwan Tso & Yoshida, 2000, syn. n. is synonymised with Nesticella 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), Nesticella songi Chen & Zhu, 2004 and Nesticella 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.


Introduction
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 (World Spider Catalog 2016) and 12 fossil species belonging to four extinct genera (Dunlop et al. 2016) are considered in this family. The majority of nesticids occur in temperate areas of the Holarctic realm where they are mainly restricted to cave-like environments and are medium-sized and long-legged with different levels of adaptation to the troglophylic life. By contrast, nesticids living in tropical or subtropical areas of the Afrotropic, Australian, Neotropic and Oriental realm are mostly characterized by being smaller than their Holarctic counterparts and they have shorter legs and only a minor bond to cave environments. They frequently occur outside caves in forest litter, on grass and under stones (Lehtinen and Saaristo 1980).
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 andGaucelmus 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 andTyphlonesticus 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 andNesticella Lehtinen &Saaristo, 1980 separated by palpal morphological characters; these genera were subsequently synonymized by Wunderlich (1986). On the contrary, Marusik and Guseinov (2003) considered the diagnosis provided by Lehtinen and Saaristo reliable, supporting the split into two distinct genera and rejecting the synonymization by Wunderlich (1986). The systematics of Nesticella remains unsolved and is a cause of debate.
Since the work of Lehtinen and Saaristo (1980), Nesticide publications have been small-scale and separate rather than large-scale taxonomic surveys. The contrasting qualitative levels of descriptions and illustrations, sometimes only basic or very schematic, made further progress on the study of this family more complex. Therefore, new species discovered in Asia or Africa, including the majority of the relatively well-studied Japanese nesticids, were all assigned to the genera Nesticella and Nesticus. Only recently has the increasing interest in spider research in China and Southeast Asia allowed a better knowledge of local species systematics and ecology. Thus, the genus Pseudonesticus was created by Liu and Li (2013a) to accommodate P. clavatus Liu & Li, 2013, a new blind troglobitic species from Yunnan, China. Three other monotypic genera, Hamus, Nescina and Wraios were erected by Ballarin and Li (2015) to include three Chinese species (H. bowoensis, N. minuta and W. longiembolus) from Tibet and Yunnan that have peculiar morphological characters. Moreover, a recent study by Zhang and Li (2013) on the Chinese Nesticella revealed that the species of the Yunnan-Guizhou Plateau represent an ancient lineage with a strong bond to the subterranean environment despite the absence of morphological adaptations. According to the authors, this troglobitic lifestyle probably evolved through multiple, independent colonization events during the Pleistocene.
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 Li 2013a, 2013b;Grall and Jäger 2016;Li and Lin 2016). However, deeper morphological and molecular studies of this material revealed a further, extraordinary species diversity in South China and in the adjacent areas that was previously widely underestimated.
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.

Material and methods
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 (Khmelik et al. 2006) and Combine ZP image stacking software. Male palps and epigynes were examined and photographed after dissection. Epigynes were treated in lactic acid before being embedded in Arabic gum to take the photos of the vulva. To reveal the course of the spermatic ducts, in some cases male palps, were also cleared using lactic acid and subsequently mounted in Hoyer's Solution. The left palp was photographed and described. All measurements are in millimeters, with leg measurements given in the following sequence: total length (femur, patella, tibia, metatarsus, and tarsus).
Abbreviations used in the text or figures are given in Table 2. References to figures in the cited papers are listed in lowercase ( fig. or figs); figures from this paper are noted with an initial capital (Fig. or Figs). All molecular vouchers are preserved in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing. Institutions hosting the type material and other specimens used in this work is reported in Table 2.

Diagnosis.
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).
Description. 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.
Diagnosis. 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 Ballarin and Li 2015: 180, fig Description. See Ballarin and Li (2015 Etymology. The specific name derives from the Latin word "cornutus" = horned, and refers to the horn-like distal process of the paracymbium; adjective. Diagnosis. 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. 1A), the sharp, hooked terminal apophysis process II (Ta-II) (Fig. 1A, D) and the horn-like distal process of the paracymbium (Dp) (Fig.  1A-B), as opposed to a long Ta-I and a needle-like, very short or caniniform Dp in H. bowoensis (see Ballarin and Li 2015: 180, fig. 1A-D) and H. mangunensis sp. n. (Fig. 5A-B, D) and as opposed to a blunt Ta-I and a pointed Dp in H. kangdingensis sp. n. (Fig. 3D, E, G). Females can be separated from those of the similar species H. bowoensis and H. luzon sp. n. by the more pear-shaped spermathecae  and the relatively straight copulatory ducts (Cd) (Fig. 2G), as opposed to the almost round spermathecae and the strongly twisted Cd in H. bowoensis (see Ballarin and Li 2015: 180, fig. 2A-C with Cd reported as Fd) and H. luzon sp. n. (Fig. 4D-F).
Description. Habitus as in Fig. 2A-D. Carapace yellowish. Mouthparts yellow in the male, light brown in the female. Sternum pale yellow in males, yellow in females. Legs uniformly pale yellow. Opisthosoma yellowish, slightly dark in males.
Male ( Etymology. The specific name is derived from the type locality; adjective. Diagnosis. Males can be easily distinguished from those of the other congeners by the caniniform distal process of the paracymbium (Dp) (Fig. 3E, H). Another character to separate them from males of Hamus bowoensis, H. cornutus sp. n. and H. mangunensis sp. n. is the wider terminal apophysis with a blunt and short process I (Ta-I) ( Fig. 3F-G), whereas the entire Ta is a large C-shape in H. bowoensis (see Ballarin and Li 2015: 180, fig. 1B-C) and H. mangunensis sp. n. (Fig. 5A, C) and shaped as an inverted comma with a missing Ta-I in H. cornutus sp. n. (Fig. 1A).
Male ( Etymology. The specific name is derived from the type locality, the island of Luzon in the Philippines; noun in apposition. Diagnosis. 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) . In the other two species, the shape of the spermathecae are more pyriform, and the fertilization ducts are longer and wider (see  4D). Spermathecae wide, strongly sclerotized, almost round and close to each other. (Fig. 4E-F). Fertilization ducts short and narrow, connected to the base of the spermathecae and located in the front of the copulatory ducts. Together they form an upturned structure ( Fig. 4E-F). Copulatory ducts short, forming a coiled structure in the center (Fig. 4F).
Female ( Diagnosis. Hamus mangunensis sp. n. is easily distinguished from the other Hamus species by a tiny dentiform distal process of the paracymbium (Dp) (Fig. 5B, D), more developed in the other species, and by the wide terminal apophysis process I (Ta-I) with a blunt tip (Fig. 5A, D) which is absent in H. cornutus sp. n. (Fig. 1A, D), very short in H. kangdingensis sp. n. (Fig. 3D, G) and narrower with a sharp point in H. bowoensis (see Ballarin and Li 2015: 180, fig. 1B-D).
Description. Habitus cannot be properly described due to the very poor condition of the sample.
Male palp (Fig. 5A-D): paracymbium weakly sclerotized, dorsal apophysis laminar and translucent, ventral apophysis broad and earlobe-shaped, distal process very tiny, sclerotized ( Fig. 5A-B, D). Terminal apophysis well-developed and strongly sclerotized, with two elongate processes (Ta-I-II) forming together a Clike structure (Fig. 5A, C, D). Conductor with two processes near its base, Cp-I short and flat, hook-like, Cp-II elongate and laminar, forming a curved groove. (Fig. 5A Diagnosis. 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.
Description. 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.
Remarks. The molecular analysis confirms that Nescina kohi sp. n. differs from the type species, N. minuta (see Appendix B). Futhermore N. kohi sp. n. is distributed in Singapore while the type locality of the other species is in Yunnan, China, with a linear distance of more than 2000 kilometers separating the two sites.

Genus Nesticella Lehtinen & Saaristo, 1980
Nesticella Lehtinen & Saaristo, 1980: 55;Wunderlich 1986: 93;Gray 1989: 88;Marusik and Guseinov 2003: 37. Type species. Nesticus nepalensis Hubert, 1973 from Nepal. Diagnosis. Nesticella belongs to the tribe Nesticellini sensu 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.
Description. 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.
Distribution. 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.
Remarks. Although the morphological characters of Nesticella species may support the hypothesis of two possible distinct genera as claimed by Lehtinen and Saaristo (1980) and Marusik and Guseinov (2003), the variability within this group is wide, and a further analyses that incorporate molecular data are required to solve the question. Therefore, we prefer to follow the current classification as given in the World Spider Catalog (2016) with all the species included in a single genus corresponding to the tribe Nesticellini sensu Lehtinen & Saaristo (1980). We propose five species-groups (brevipes, mogera, nepalensis, phami and quelpartensis-groups) based on male palpal and epigynal morphology and preliminary results of our molecular analysis of the entire Nesticidae. Two species-groups already established by Lehtinen and Saaristo (1980) (brevipes-group and mogera-group) are followed while the nepalensis-group coincides with the genus Nesticella sensu Lehtinen & Saaristo (1980). Two more new groups (phami-groups and quelpartensis-group) are here established.
Incertae sedis. Two recently-described species from Laos and Myanmar, Nesticella foelixi Grall &Jäger, 2016 andN. 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.

Nesticella brevipes-group
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.
Description. Habitus as in Fig. 9A-D. Carapace uniformly pale yellow in males, yellowish with a "V"-shaped dark mark on the cephalic area in females. Ocular area with a long seta between AMEs. Cervical groove and fovea indistinct. Mouthparts brown yellow. Sternum orange in males, faintly dark in females, with sparse long setae in the center, thin and short near the margins. Legs and female palps yellowish, distally darker. Opisthosoma yellowish with faint dark areas.
Male palp ( Male ( Diagnosis. 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. 15A-F), by the vestigial eyes (Fig. 11A, 11C); further differences can be found in the shape of the ventral (Va) and distal (Dp) processes of the paracymbium ( Fig. 10A-B, D) and in the epigynal scape. Nesticella caeca sp. n. can be separated from N. gazuida sp. n. by the different configuration of the fertilization (Fd) and copulatory ducts (Cd) (straighter, thicker and shorter in N. caeca sp. n. than in N. gazuida sp. n.) (Fig. 11G vs. Fig. 15F).
Description. Habitus as in Fig. 11A-D. Carapace pale yellow. Six vestigial eyes, AME absent, the others strongly reduced. Cervical groove and fovea indistinct. Mouthparts pale yellow. Sternum pale yellowish with long setae. Legs with the same color of the carapace but slightly darker. Opisthosoma uniformly greyish with faint darker areas.
Remarks. The molecular analysis shows that there are certain differences between Nesticella caeca sp. n. and the close related N. gazuida sp. n. (see Appendix B). Etymology. The specific name is derived from the type locality, the city of Chongqing; noun in apposition.
Diagnosis. The new species is closely related to Nesticella odonta (see Figs 24A-D, 25A-G and Chen and Zhang 1991: 158, figs 1-6) and N. xiongmao sp. n. (Figs 35A-D, 36A-G). It can be separated from the former species by the more protruding and almost round scape (Sp) (Fig. 12D vs. Fig. 25E) and the narrower copulatory ducts (Cd) (Fig. 12F vs. Fig. 25G); it can be distinguished from the latter species by the less protruding, wider scape ( Fig. 12D vs. Fig. 36E) and the larger distance between the copulatory ducts (Cd) (Fig. 12F vs. Fig. 36G). The short and slightly round scape and the straight and almost parallel copulatory (Cd) ducts allow the separaton of N. chongqing sp. n. from all the other species of the brevipes-group.
Description. Habitus as in Fig. 12A-C. Carapace yellow with wide dark areas. Sternum dark grey. Cephalic area with several long setae around the ocular area and along the midline. Cervical groove and fovea distinct. Mouthparts yellow. Sternum with sparse black setae. Legs and palps proximally yellowish, distally darker with dark rings around each tibia and metatarsus. Opisthosoma with a light background and pairs of dark areas partially merged into each other on the dorsal, lateral and ventral sides forming a light, double cross-like mark. Epigyne ( Fig. 12D-F): dark with a lighter mark in the center. Scape short, approximately two to three times wider than long. Posterior and lateral margin weakly sclerotized, slightly protruding beyond the epigynal posterior margin (Fig. 12D). Spermathecae nearly ovoid, separated by about 1.5 diameters (Fig. 12F). Fertilization ducts thin and long, reaching the spermathecae with 3.5 coils. Copulatory ducts short, slightly narrower at their distal end ( Fig. 12E-F).  Etymology. The specific name is derived from the type locality; adjective. Diagnosis. 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. 13A-B); by the thin and sharp tegular apophysis (Tg) (Fig. 13A) and by the narrower cymbium when observed in dorsal view (Fig. 13B). Diagnostic characters for the females are the remarkably wide spermathecae (S) (Fig. 14F) and the short, fertilization (Fd) and copulatory (Cd) ducts each with only a few coils (Fig.  14G) which are respectively smaller and longer/more coiled in the other species of the same group.
Description. Habitus as in Fig. 14A-D. Carapace yellow, dark at margins and in the midline. Clypeal area yellowish. A single long seta present between AMEs. Cervical groove faintly visible, fovea distinct. Mouthparts orange. Sternum faintly dark, with sparse long setae. Legs and female palps yellowish, distally darker in each segment. Opisthosoma with a yellowish background and distinct dark pattern formed by transverse bands. Wide dark band between the epigastral furrow and spinnerets.
Male palp (    Etymology. The specific name is derived from the name of the cave where the species was collected; noun in apposition. Diagnosis. 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. 11A-G), by their pale coloration and the reduced eyes (Fig. 15A). N. gazuida sp. n. can be separated from N. caeca sp. n. by the smaller spermathecae and especially by the longer fertilization (Fd) and copulatory (Cd) ducts, clearly bent in the middle, which are shorter, more compact and straighter in the latter species (Fig. 15F vs. Fig. 11G).
Female ( Etymology. The specific name is derived from the type locality; adjective. Diagnosis. 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. 16B), a rectangular distal process on the paracymbium (Dp-I) ( Fig. 16D) and, for the females, by the appearance of the short scape ( Fig. 17E) and the thick copulatory ducts ( Fig. 17F-G). Males can be separated from those of N. semicircularis (see Liu and Li 2013b: 521, figs 19-22) by the dark pattern, the wider dorsal apophysis (Da), by the wider distal process (Dp-I), and the stockier tegular apophysis (Tg) (Fig. 16A, 16D vs. figs 20A, 21A-C). Females of N. hongheensis sp. n. are recognized by the black marks on the opisthosoma, the wider and much shorter scape and the shape of the copulatory ducts (Cd), strongly bent in the middle ( Description. Habitus as in Fig. 17A-D. Carapace pale yellow in males, darker in females; dark in the midline, at margins and at the cephalic area. Cephalic area with several setae on the clypeus and along the cervical groove. Cervical groove and fovea distinct. Mouthparts pale yellow. Sternum greysh in males, darker in females. Legs and female palps yellowish, distally darker in each segment. Opisthosoma black with a light mark followed by three pairs of spots on the dorsal and lateral sides. The whole pattern is more evident in females.
Male palp ( Fig. 16A-D): cymbial surface finely reticulated, densely covered with thick, long setae (Fig. 16B). Paracymbium with the first ventral process wide with a sharp tip, the second smaller and blunt (Fig. 16A, D). Two compact distal processes laterally compressed, Dp-I wide and strongly sclerotized, Dp-II short and stocky. Dorsal apophysis triangular, broad and flat (Fig. 16B). Terminal apophysis blunt, translucent and fingerlike (Fig. 16A, C). Tegular apophysis short and stocky (Fig. 16A). Conductor with a thin process (    Etymology. The species is named after the Jingpo people, an ethnic minority living in the Gaoligongshan region of Yunnan Province; noun in apposition. Diagnosis. This new species is closely related to N. lisu sp. n. (Fig. 20F-G), but can be distinguished from the latter species by the subglobular spermathecae (S) (piriform in N. lisu sp. n.), the more compact and less coiled fertilization ducts (Fd) and the wider copulatory openings (Co) (Fig. 18E-F vs. Fig. 20F-G). The strongly coiled fertilization ducts (Fd) and the general shape of scape (Sp) and copulatory ducts (Cd) allow the separation of N. jingpo sp. n. from all other species of the brevipes-group.
Description. Habitus as in Fig. 18A-C. Carapace pale yellow, darker near the center and the margins. Ocular area with several setae. Cervical groove and fovea distinct. Mouthparts pale yellow. Sternum dark greyish, bearing sparse setae. Legs and female palps yellowish, distally darker in metatarsi and tarsi. Opisthosoma with four pairs of pigmented marks on the dorsal side, the first pair extended to laterals. Ventral side with a wide median band. Epigyne ( Fig. 18D-F): reddish brown surrounded by an extensive dark area (Fig.  18D). Scape short, translucent, nearly as wide as the diameter of the spermathecae (Fig. 18E). Spermathecae ovoid, separated from each other by about 1.5 diameters ( Fig. 18E-F). Fertilization ducts thin and long, coiling into 4.5 loops before reaching the spermathecae (Fig. 18E  Male ( Etymology. The specific name is derived from the type locality; adjective. Diagnosis. 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. 30E-G) and N. machadoi from Angola (see Hubert, 1971: 75, figs 1-5) are morphologically similar to N. liuzhaiensis sp. n. The new species can be separated from N. robusta sp. n. by the straighter and less tortuous copulatory ducts (Cd) (Fig. 21F vs. Fig. 30G) and from N. machadoi by the smaller spermathecae (S), the more widely separated copulatory ducts (Cd) and the straight rather than concave posterior margin of the scape (Fig. 21G vs. fig. 3). Furthermore, in N. liuzhaiensis sp. n., the PME are equal to ALE in size while the PME are larger than the ALE in N. machadoi (Fig. 21A vs. fig. 1).
Description. Habitus as in Fig. 21A-C. Carapace dark yellow, with extended dark areas near the center and the margins. Cephalic area with several long setae at the midline and along the cervical groove. Cervical groove and fovea distinct. Thoracic area with faint radial furrows. Mouthparts brown-yellowish. Sternum dark, with long setae. Legs and female palps yellowish, tarsus distally darker. Opisthosoma light yellow with paired dark spots on the dorsal side, often merged with each other and forming a lighter cross-shaped mark on the background. Epigyne (

Remarks.
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 Lehtinen and Saaristo (1980) on the basis of a single female; the specimens have the very short and wide, squared scape, rarely seen in the Nesticella species of the brevipes-group. The small, almost round spermathecae and the long copulatory ducts that are slightly curved outward and visible through the tegument are also the same. Although described from two different countries, the type localities are close to the common border between Myanmar and Thailand and are approximately 200 km apart. Based on the evidence mentioned above, we propose the synonymy of N. inthanoni Lehtinen & Saaristo, 1980with N. mollicula (Thorell, 1898 Etymology. The specific name is derived from the type locality; adjective. Diagnosis. 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. 22A vs. Fig. 8A) and by the larger scape (Sp) and the wider space between the spermathecae (S) (Fig. 23G vs. Fig. 9G). It is recognized from N. songi by the wider triangular tegular apophysis (Tg) bearing a sharper tip, the smaller process of the conductor (Cp), the shorter process I of the ventral apophysis (Va-I), the more pointed ventral process II (Va-II) (Fig. 22A, D vs. Fig. 33A, D), the wider distance between the spermathecae (S) and the ducts (Cd and Fd) oriented outward rather than almost parallel (Fig. 23G vs. Fig. 34G). The new species can be distinguished from N. verticalis (see Liu and Li 2013b: 522, figs 27-30) by the sharper tegular apophysis (Tg), the wider and more squared distal process I of the paracymbium (Dp-I) when observed dorsally, the shorter and more compact copulatory ducts (Cd), the shorter scape (Sp) and the spermathecae (S) being more oriented outward (Figs 22A, D, 23G vs. figs 27B, 29B, D, 28A, D, 30C). The absence of a dorsal apophysis (Da), the different shape of  the distal processes of the paracymbim (Dp) (especially the Dp-I) and the shape of the tegular apophysis (Tg) help to distinguish males of N. nandanensis sp. n. from those of all the other species of the group.
Description. Habitus as in Fig. 23A-D. Carapace pale yellow to beige, with faint dark areas around the cervical furrow, the fovea and near the margins. Cervical groove and fovea distinct. Mouthparts yellow. Sternum yellowish with sparse setae. Leg uniformly yellowish, distally darker tibiae, metatarsi and tarsi. Opisthosoma pale greyish, faintly darker on the dorsal and posterior side and around the spinnerets.
Male ( Diagnosis. This species can be easily recognized from the others belonging to the brevipes-group, with the exception of N. xiongmao sp. n. (Fig. 35A-  bium (Dp-I) that is bent downward (Fig. 24A, D), the lobed distal process II (Dp-II) (Fig. 24D) and the sharp tegular apophysis (Tg) (Fig. 24A, C-D). Females can be distinguished by the short scape (Sp) with a flat posterior margin (Fig. 25E) and by the compact, almost straight ducts (Fd and Cd) (Fig. 25F-G). Males of N. odonta can be separated from those of N. xiongmao sp. n. by the wider distal process I of the paracymbium (Dp-I) (Fig. 24A vs. Fig. 35A) and the broader and more lobed distal process II (Dp-II) (Fig. 24D vs. Fig. 35D). Females are distinguished by the shorter, squared scape (Sp) with a flat rather than rounded posterior margin (Fig. 25E vs. Fig. 36E), the folded rather than straight copulatory ducts (Cd) and the greater distance between the spermathecae (S) (Fig. 25F-G vs. Fig. 36F-G). The same combination of characters allows females of N. odonta to be distinguished from those of N. chongqing sp. n.
Remarks. 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. 24A-D and Tso and Yoshida 2000: 14, figs 4-6). Furthermore, N. odonta seems to be a widely distributed species in throughout southern China, and its presence in Taiwan is very likely. In addition, the in progress morphological and molecular analysis of newly collected male and female specimens from the type locality of N. taiwan demonstrate that the females illustrated by Tso & Yoshida as N. taiwan were mismatched, belonging instead to a different species, N. kaohsiungensis sp. n.. Based on the above mentioned evidence, we propose the synonymy of N. taiwan Tso & Yoshida, 2000 with N. odonta (Chen, 1984 Etymology. The specific name is derived from the type locality; adjective. Diagnosis. This new species is similar to Nesticella gracilenta (see Liu and Li 2013b: 512, figs 13-16), N. xixia sp. n. (Figs 37A-D, 38E-G), N. semicircularis (see Liu and Li 2013b: 521, figs 19-22) and N. shanlinensis (see Liu and Li 2013b: 521-522, figs 23-26) for the general shape of vulva. It can be separated from the first three species by the wider and longer scape (Sp) (Fig. 26D), the longer and straighter ducts (Fd and Cd) (Fig. 26F), the more coiled fertilization ducts (Fd) (Fig. 26F) and the darker carapace (Fig. 26A, C). N. qiaoqiiensis sp. n. can be distinguished from N. shanlinensis by the longer ducts provided with more coils (Fig. 26F vs. figs 24C, 26D). The wide, lobed scape, the short copulatory ducts and the general shape of the strongly coiled fertilization ducts (Fd) (Fig.  26E-F) easily separate the new species from all other species of the brevipes-group.
Description. Habitus as in Fig. 26A-C. Carapace pale yellow, dark at margins and in the center. Cervical groove and fovea indistinct. Mouthparts yellowish brown. Sternum yellow, strongly pigmented, with sparse long setae. Female palps and legs yellow, proximally lighter in femora. Opisthosoma greyish with irregular dark marks.
Epigyne (Fig. 26D-F): weakly sclerotized, translucent. Reddish brown vulva (Fig.  26D). Scape short, flat and lobed, with almost round margins, translucent, laterally sclerotized, protruding beyond the epigynal posterior margin (Fig. 26D). Spermathecae ovate, separated by about 1.6 diameters (Fig. 26F). Fertilization ducts long and strongly coiled, reaching the spermathecae with four loops. Copulatory ducts short, mesially swollen and basally narrower ( Fig. 26E Etymology. 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. Diagnosis. 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.  27B) and the stocky, hooked distal process I (Dp-I) (Fig. 27D) in the males; by the wide and short scape (Sp) with a convex posterior margin and the almost straight ducts (Fd and Cd) in the females (Fig. 28G). It can be differentiated from N. falcata (see Liu and Li 2013b: 511, figs 9-12) by the shorter and blunt distal process II (Dp-II), the stockier distal process I (Dp-I) (  N. liuzhaiensis sp. n.) and by the shorter distance between the spermathecae and the copulatory ducts (Fig. 28G vs. Fig. 21F). It can be distinguished from N. robusta sp. n. by the thicker distal process I (Dp-I), the longer ventral process II (Va-II), strongly reduced in N. robusta sp. n. (Fig. 27A-B, D vs. Fig. 29A-B, D), and by the straighter copulatory ducts (Fig. 28G vs. Fig. 30G).
Description. Habitus as in Fig. 28A-D. Carapace pale yellow in males, darker in females, with a dark color around the margins and at the cephalic area. Thoracic area marginally pigmented. Several sparse setae present at the ocular area and along the cervical groove. Cervical groove distinct. Fovea deep. Mouthparts pale yellow in the male, darker in the female. Sternum lighter in the male than in the female. Legs and female palps yellowish, distally darker in tibiae, metatarsi and tarsi. Opisthosoma yellowish with paired black marks on the dorsal side, partially merged each other; the dark spots are extended to the lateral and ventral sides. Darker and more evident pattern in females. Spinnerets and colulus pale yellow in males, darker in females.
Male ( Etymology. 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.
Diagnosis. Males of Nesticella robusta sp. n. can be easily recognized by the single,     Liu and Li 2013b: 521, figs 19-22). It can be easily separated from the first species by the presence of fully developed eyes (Fig. 32C vs. Fig. 15A), the wider spermathecae (S) and the thicker ducts (Fd and Cd) (Fig. 32G vs. Fig. 15F). Males can be recognized from those of N. xixia sp. n. by the much wider ventral process I (Va-I), the shorter and thicker distal process I (Dp-I), the blunter tegular apophysis (Tg) (Fig. 31A, D vs. Fig. 37A, D); females are distinguished by the straight posterior margin and the longer and slimmer ducts (Fd and Cd) (Fig. 32G vs. Fig. 38G). N. sanchaheensis sp. n. can be distinguished from N. semicircularis by the wider tegular apophysis (Tg), the wider ventral process I (Va-I), the different shape of the distal process I (Dp-I) when viewed dorsally ( Fig. 31A Description. Habitus as in Fig. 32A-D. Carapace pale yellow. Cervical groove faint, fovea indistinct. Mouthparts darker than the carapace. Sternum yellowish covered with sparse long setae. Legs and female palps yellowish, distally darker in each tibiae, metatarsi and tarsi. Opisthosoma uniformly grey, darker in females. Male palp (Fig. 31A-D): paracymbium with two wide ventral processes, Va-I flat, wide and sharp, Va-II short, narrow and blunt (Fig. 31A, D); distal process strongly sclerotized with two branches (Fig. 31B), Dp-I small and stumpy, Dp-II bigger and pointed. Triangular dorsal apophysis laminar and wide (Fig. 31B). Terminal apophysis blunt, horn-like, translucent, and with a granulated surface. Tegular apophysis triangular, weakly sclerotized (Fig. 31A). Conductor ending with a short, spout-shaped, sclerotized process (Fig. 31A, C-D).
Male ( Diagnosis. 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. 33A vs. Fig. 22A), by the narrower distal process I of the paracymbium (Dp-I) in dorsal view (Fig. 33B vs. Fig. 22B), by the shorter ventral process I (Va-I) and the blunter ventral process II (Va-II) (Fig. 33A, D vs. Fig. 22A, D). They can be separated from males of N. baiseensis sp. n. by the shorter and thicker tegular apophysis (Tg), by the more squared distal process I (Dp-I), by the longer ventral process I (Va-I) and the thicker ventral process II (Va-II) ( Fig. 33A-B, D vs. Fig. 8A-B, D). Females of N. songi can be recognized from those of N. nandanensis sp. n. by the narrower space between the spermathecae (S) and the more parallel ducts (Fd and Cd) rather than ducts that are oriented outward (Fig. 34F-G vs. Fig. 23F-G). They are distinguished from females of N. baiseensis sp. n. by the wider scape (Sp) and the narrower and more twisted ducts (Fd and Cd) (Fig. 34G vs. Fig. 9G). Females of N. songi are distinguished from those of N. yao sp. n. by the almost round scape (Sp) with a convex and smooth posterior margin rather than an almost flat and wrinkled scape, by less straight ducts (Fd and Cd) (Fig. 34E-G vs. Fig. 39D-F).
Description. See Figs 33A-D, 34A-G and Chen and Zhu (2004). Habitat. Cave. Distribution. China (Guizhou) (Fig. 82). Etymology. 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.
Diagnosis. The new species can be easily distinguished from all the species belonging to the brevipes-group, with the exception of Nesticella odonta and N. chong-    qing sp. n., by the sharp, hook-like distal process I of the paracymbium (Dp-I) that is bent downward (Fig. 35D), the lobed distal process II (Dp-II) (Fig. 35D) and the triangular tegular apophysis (Tg) with a sharp tip (Fig. 35A). Females can be distinguished by the tongue-like, protruding scape (Sp) with an almost round posterior margin (Fig. 36F) and by the compact, straight ducts that are almost parallel to each other (Fig. 36G). N. xiongmao sp. n. can be recognized from N. odonta by the thinner distal process I of the paracymbium (Dp-I) (Fig. 35A, D vs. Fig. 24A, D) and the shorter and less lobed distal process II (Dp-II) (Fig. 35D vs. Fig. 24D); females are distinguished by the longer, tongue-like scape (Sp) with an almost round posterior margin rather than a flat scape (Fig. 36E-F vs. Fig. 25E-F), by the more compact and straighter ducts (Fd and Cd) and by the shorter distance between the spermathecae (S) (Fig. 36F-G vs. Fig. 25F-G). Females of N. xiongmao sp. n. can be separated from those of N. chongqing sp. n. by the more protruding and narrower scape and by the narrower space between the copulatory ducts (Cd) (Fig. 36F-G vs. Fig. 12E-F).
Description. Habitus as in Fig. 36A-D. Carapace pale yellow, dark at margins and near the cephalic area; darker in females. Ocular area dark. Cervical groove and fovea distinct. Mouthparts brown-yellowish. Sternum pigmented, darker in female. Legs and female palps yellowish, distally darker and with a dark ring at each tibia and metatarsus. Opisthosoma covered with long setae, greyish, with paired black marks on the dorsal, lateral and ventral sides, partially fused to each other. The whole pattern is more evident in males.
Epigyne (Fig. 36E-G): greyish. Scape lobed and protruding out of the epigynal posterior margin, tongue-like, about two times longer than wide, and showing an almost round posterior margin (Fig. 36F). Spermathecae small, ovate, separated each other by about 1.3 diameters (Fig. 36F-G). Ducts compact and straight, almost parallel to each other. Fertilization ducts thick and long, reaching the spermathecae with 2.5 loops (Fig. 36G). Copulatory ducts thick and short (Fig. 36F).
Male ( Habitat. Forest leaf litter. Distribution. Known only from the type locality (Fig. 82). Diagnosis. 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. 37A, D vs. Fig. 31A, D); females are separated by the shorter ducts (Fd and Cd), the smaller spermathecae (S) and the almost round posterior margin of the scape (Sp) (Fig. 38G vs. Fig. 32G) which is flat in N. sanchaheensis sp. n.. N. xixia sp. n. can be distinguished from N. semicircularis (see Liu and Li 2013b: 521, figs 19-22) by the slimmer ventral process I of the paracymbium (Va-I), the sharper tegular apophysis (Tg) and the different shape of the distal processes (Dp-I, Dp-II) (Fig. 37A, D vs. figs 19A, 20A-B, 21A-B, D) for the males; females instead are recognized by the wider scape (Sp), the smaller spermathecae (S) and the larger distance between the copulatory ducts (Cd) (Fig. 38E-G vs. figs 20C-E, 22C-E). The general shape of the ventral and distal processes of the paracymbium (Dp-I and Dp-II), the short copulatory ducts (Cd) and the lobed scape (Sp) allow easy separation from all the other species of the brevipes-group.
Description. Habitus as in Fig. 38A-D. Carapace pale yellow. Cervical groove faint, fovea distinct. Mouthparts slightly darker than the carapace. Legs uniformly pale yellow. Female palpal femur and patella yellowish, tibia and tarsus darker yellow. Opisthosoma uniformly greyish with faint dark marks. Dorsum darker than venter.
Male ( Fig. 34E-G, and Chen & Zhu, 2004: 87, figs 6-7) and N. nandanensis sp. n. (see Fig. 23E-G). It can be separated from the former species by the squarer scape (Sp) with a flat and furrowed posterior margin, by the straighter and more compact ducts (Fd and Cd) and by the lack of a clear constriction around the coils of the fertilization ducts (Fd) (Fig. 39D-F vs. Fig. 34E-G). It can be recognized from N. nandanensis sp. n. by the squarer scape (Sp) with a flat rather than convex posterior margin, by the straighter copulatory ducts (Cd) (bent outward in N. nandanensis sp. n.) and the spermathecae (S) closer to each other (Fig. 39D-F vs. Fig. 23E-G).

Nesticella mogera-group
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.  Fig. 47A, D). Females can be separated from those of N. huomachongensis sp. n. by the different shape of the posterior margin of scape (convex in N. fuliangensis sp. n., straight in N. huomachongensis sp. n.) (Fig. 41E-G vs. Fig. 43E-G); from those of N. yanbeiensis sp. n. by the shorter and wider scape (Sp) and by the greater distance between the copulatory ducts (Cd) (Fig. 41E-G vs. Figs 48E-G). The same combination of characters allows an easy separation from all the other species of the group.
Male ( (Fig. 42B vs. Fig. 45B), the shorter process of the conductor (Cp) (Fig.  42D vs. Fig. 45D) and the different shape of the distal process of the paracymbium (Dp) which appears stockier in N. wanzaiensis sp. n. (Fig. 42A vs. Fig. 45A). The wider process of the conductor (Cp) and the more squared distal process of the paracymbium (Dp) (Fig. 42A, D vs. Fig. 40A, D) allow the separation of males of the new species from those of N. fuliangensis sp. n. Females can be recognized by the wider and squarer scape (Sp) (narrower in N. wanzaiensis sp. n.) and by the truncate posterior margin (curved in N. fuliangensis sp. n.) (Fig. 43F-G vs. Fig. 46F-G vs. Fig. 41F-G). The same combination of characters allows easy separation of N. huomachongensis sp. n. from the other species of the mogera-group.
Description. Habitus as in Fig. 43A-D. Carapace yellowish. Ocular area slightly protruding in males. Cervical groove and fovea indistinct. Mouthparts yellowish, darker in females than in males. Sternum pale yellow. Legs uniformly yellowish. Female    palpal femur pale yellow, tibia and tarsus brown-yellowish. Opisthosoma grey in males, grey-yellowish in females.
Description. See Marusik and Guseinov (2003 Etymology. The specific name is derived from the type locality; adjective. Diagnosis. 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. 44E,  G). Furthermore, the new species can be recognized by the shorter and stockier distal process of the paracymbium (Dp), by the different shape of the process of the conductor (Cp) and the terminal apophysis (Ta) and by the narrower ventral apophysis (Va) (Fig. 44A-D).
Description. Habitus as in Fig. 44E-G. Carapace pale yellow, with sparse short setae. Ocular area with few long setae, eyes completely lacking (Fig. 44E, G). Cervical furrow and fovea indistinct. Mouthparts pale yellow. Sternum pale yellow. Legs uniformly pale yellowish. Opisthosoma grey with long setae.
Male palp (Fig. 44A-D): paracymbium with a sharp, straight, ventral apophysis and a blunt, strongly sclerotized distal process ( Fig. 44A-B, D). Terminal apophysis well-developed with granulate surface (Fig. 44A). Conductor ending with a strongly sclerotized, finger-like process (Fig. 44A, C-D     45A, D vs. Fig. 42A, D). N. wanzaiensis sp. n. can be easily distinguished from N. fuliangensis sp. n. by the stockier and thicker process of the conductor (Cp) (Fig. 45A, D vs. Fig. 40A, D). Females are distinguished by the narrower and protruding scape, which is shorter and stockier in females of the other two species, and by the narrower part of where the fertilization ducts (Fd) coils reach the spermathecae, much slimmer than the copulatory ducts (Cd) (almost of the same diameter in females of the other species) (Fig. 46E-G vs. Fig. 43E-G vs. Fig. 41E-G).
Description. Habitus as in Fig. 46A-D. Carapace yellowish. Cervical furrow and fovea indistinct. Mouthparts uniformly yellowish, darker in females. Sternum yellowish. Legs uniformly pale yellow. Opisthosoma uniformly grey in males, with a longitudinal lighter strip and two pairs of black dots in females.
Epigyne (Fig. 46E-G): posterior margin of the scape weakly sclerotized (Fig. 46E). Scape short and narrow, lightly protruding out of the epigynal posterior margin, about as wide as the diameter of the spermathecae (Fig. 46E). Fertilization ducts short and swollen, reaching the spermathecae with two loops; this area considerably narrower than the copulatory ducts (Fig. 46G). Copulatory ducts short and swollen, strongly sclerotized (Fig. 46G). Spermathecae small and globular, as wide as the copulatory ducts, separated by about two diameters.
Male ( Etymology. The specific name is derived from the type locality; adjective. Diagnosis. 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. 47A, D); diagnostic characters for the females are the short, lightly protruding scape (Sp) and the copulatory ducts (Cd) close to each other (Fig. 48E-G). N. fuliangensis sp. n. (see Figs 40A-D, 41A-G) seems to be more closely related to N. yangbeiensis sp. n., but the males can be distinguished by the thicker and more undulate process of the conductor (Cp) and by the wider and rectangular distal process of the paracymbium (Dp) (slimmer in males of N. fuliangensis sp. n.) (Fig. 47A, D vs. Fig. 40A, D). Females of the new species can be recognized by the relatively narrower and more protruding scape (Sp) and by the copulatory ducts which are closer to each other (Cd) (Fig. 48E-G vs. Fig. 41E-G).
Description. Habitus as in Fig. 48A-D. Carapace pale yellow in the male, yellow in females. Cervical groove and fovea distinct. Mouthparts brown-yellowish in males, brownish in females. Sternum pale in males, yellowish in females. Legs uniformly pale yellow. Female tibia and tarsus brown. Opisthosoma greyish, covered with long setae, with four faint greyish dorsal marks. Epigastric area pigmented in males.

Nesticella nepalensis-group
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 con-ductor (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 (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, andN. zhiyuani  Etymology. 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. Diagnosis. The new species is similar to Nesticella benoiti (see Hubert 1970: 364, figs 5-8) and N. griswoldi sp. n. (Figs 53A-D, 54A-B), although the barcode can be used to separate the latter species (see Appendix B). It can be distinguished from N. benoiti by the longer terminal apophysis (Ta), the absence of a clearly serrated tegular apophysis (Tg) and the different shape of the distal processes of the paracymbium (Dp-I, Dp-II) (Fig. 49C-D vs. fig. 6). Nesticella baobab sp. n. can be distinguished from N. griswoldi sp. n. by the slightly protruding process II of the ventral apophysis (Va-II) and the slightly shorter distal process II (Dp-II) (Fig. 49C-D vs. Fig. 53C-D). The same combination of characters allows separation of this species from all the others of the nepalensis-group.
Male ( Distribution. Known only from the type locality (Fig. 83). Wunderlich, 1995 Figs 50, 51, 83 Nesticella connectens Wunderlich, 1995: 568, figs 40-42 (♂).  (Tg-II) which is reduced in males of the other two species, by the longer and sharper terminal apophysis (Ta) and by the different shape of the ventral process I (Va-I) which is more bent in males of N. tibetana sp. n. and slimmer in those of N. nepalensis (Fig. 50A, C-D vs. Fig. 59A, 59C-D vs. Fig. 55A, C-D). Females can be separated from those of N. tibetana sp. n. by the less coiled fertilization ducts (Fd) and the longer copulatory ducts (Cd) (Fig. 51G vs. Fig. 60G), from those of N. nepalensis by the more prominent scape (Sp) (Fig. 51G vs. Fig. 56G) and from those of N. potala sp. n. by the more convoluted copulatory ducts (Cd) and the rounder spermathecae (S) (Fig.  51G vs. Fig. 57G).

Nesticella connectens
Description. Habitus as in Fig. 51A-D. Carapace yellow, faintly pigmented around the radial furrows and the margins. Cervical groove distinct. Fovea shallow. Thoracic area with distinct radial furrows. Mouthparts yellow in males, brown-yellowish in females. Sternum light yellow, with sparse long setae. Legs and female palps yellowish, distally darker in tibiae, metatarsi and tarsi. Opisthosoma light yellow with dark marks in the posterior half and around the spinnerets, darker pigmentation in males.  Male palp (Fig. 50A-D): paracymbium strongly sclerotized, Va-I elongate, ending with an almost round tip, Va-II short and rectangular; bifurcated distal process with two branches, Dp-I blunt, stocky and swollen, Dp-II elongate and sharper ( Fig.  50A-B, D). Terminal apophysis long and sharp, wrinkled, basally wide and distally narrow and curved, located behind the process of the conductor (Fig. 50A, C). Tegular apophysis sclerotized and nodular, secondary tegular apophysis well developed, protruding outward from the tegulum. (Fig. 50A, C-D). Conductor with a sclerotized, short, beak-shaped process (Fig. 50A).
Epigyne (Fig. 51E-G): weakly wrinkled and faintly pigmented. Scape square, about as wide as long and weakly sclerotized (Fig. 51E-F). Spermathecae nearly globular, separated by about 2/3 of their diameter (Fig. 51F-G). Fertilization ducts long, reaching the spermathecae with two loops (Fig. 51G). Copulatory ducts thick and straight, distally as wide as the spermathecal diameter (Fig. 51F-G Habitat. Forest leaf litter, cave. Distribution. Malaysia, Thailand (first record) (Fig. 83). Remarks. Nesticella connectens was described by Wunderlich (1995) from Peninsular Malaysia based only on males. The collection of both sexes on the Thai side of the Malay Peninsula (approximately 300 km north of the type locality) allows a detailed analysis of this species and the description of the male for the first time. Etymology. The specific name is derived from the type locality; adjective. Diagnosis. This new species is closely related to Nesticella yui (see Fig. 62A-F), but can be distinguished from the latter by the narrower scape (Sp), the narrower space between the copulatory openings (Co) and the different shape of copulatory ducts (Cd) (Fig. 52F vs. Fig. 62F). The small scape (Sp), only faintly visible, and the V-shaped copulatory ducts (Cd) allow the separation of this species from all the others belonging to the nepalensis-group.
Description. Habitus as in Fig. 52A-C. Carapace yellow, faintly pigmented in the cephalic area and at margins. Cervical groove and fovea distinct. Mouthparts yellowish. Sternum yellow with sparse long setae and additional thin setae at margins. Legs and female palps yellowish, distally darker in metatarsi and tarsi. Opisthosoma pale yellowish, and with paired dark marks, partially connected each other in the dorsal area, bordering a light, cross-shaped mark in center. Other dark markings present laterally, at the epigastric and hypogastric area and around the spinnerets. Epigyne (Fig. 52D-F): wrinkled and translucent, scape strongly reduced and barely visible (Fig. 52D-F). Copulatory openings close to each other near the posteromargin of the epigyne (Fig. 52F-G). Spermathecae almost round, separated by about 1.3 diameters. Fertilization ducts thin and long, coiled into 2.5 loops until reaching the spermathecae (Fig. 52F). Copulatory ducts thick (Fig. 52E-F Diagnosis. The new species is similar to Nesticella benoiti (see Hubert 1970: 364, figs 5-8), N. baobab sp. n. (see Fig. 49A-G) and N. vanlang sp. n. (see Fig. 61A-F). Males can be distinguished from those of N. benoiti by the longer terminal apophysis (Ta), the shorter tegular apophysis (Tg) lack of a deep serration, and by the longer, sharper distal process II of the paracymbium (Dp-II) (Fig. 53A-B, D vs. fig. 6). Males of N. griswoldi sp. n. can be separated from those of N. baobab sp. n. by the slightly wider process II of the ventral apophysis (Va-II) and the slightly longer distal process II (Dp-II) (Fig. 53A-B, D vs. Fig. 49A-B, D). Females can be recognized from those of N. benoiti and N. vanlang sp. n. by the short, rectangular scape (Sp) (narrower and pointed in N. benoiti, shorter and less evident in N. vanlang sp. n.) and by the distance between the spermathecae (S) (close to each other in N. vanlang sp. n.) (Fig. 54E-G vs. figs 7-8 vs. Fig. 61D-F). The same combination of characters allows separating this species from all the others belonging to the nepalensis-group.
Description. Habitus as in Fig. 54A-D. Carapace pale yellow in males, yellow in females. Cervical groove and fovea distinct. Mouthparts yellow. Sternum yellow, dark-ish pigmented in males. Legs and female palps pale yellow, distally darker in metatarsi and tarsi. Opisthosoma pale yellow, with faint grey dark and covered with long setae.
Male ( Habitat. Rain forest leaf litter. Distribution. Known only from the type locality (Fig. 83).
Habitat. Forest leaf litter, under stone. Distribution. China (Tibet) (first record) (Fig. 83) Etymology. 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.
Diagnosis. The new species is closely related to Nesticella tibetana sp. n. (see Fig.  60C-G), N. nepalensis (see Fig. 56C-G) and N. connectens (see Fig. 51C-G). It can be distinguished by the triangular spermathecae (S), the 45° angle made by the copulatory ducts (Cd) (approx. 90° in the other species), the short fertilization ducts lacking coils, and the lack of a clear abdominal pattern (Fig. 57A-F). All these charaters have no similarities to all the other species of the nepalensis-group.
Description. Habitus as in Fig. 57A-C. Carapace yellow, with short setae near the clypeus and the thoracic area, long setae near the cephalic midline and cervical groove. Cervical groove distinct, fovea shallow. Mouthparts brown-yellowish. Sternum flat, faintly reticulated in the center, with both long and short setae at its margins. Legs and female palps yellowish, distally darker in metatarsi and tarsi. Opisthosoma greyish and faintly pigmented with darkish spots. Spinnerets yellow, colulus yellowish. Epigyne (Fig. 57D-F): wrinkled and translucent (Fig. 57D). Scape short and rectangular, approximately two times wider than long (Fig. 57E-F). Spermathecae triangular, separated by about their length. Fertilization ducts thin, without complicated coils and only one loop before reaching the spermathecae (Fig. 57F). Copulatory ducts thick and straight, laterally bent in the middle (Fig. 57E-F).

Habitat. Forest leaf litter.
Distribution. Known only from the type locality (Fig. 83). Etymology. The specific name is derived from the Island of Sulawesi were this species was collected; noun in apposition.
Diagnosis. This new species can be distinguished from all the others of the nepalensisgroup 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. 58E-F). It differs from N. yui (see Fig. 62A-F; Wunderlich & Song, 1995: 347, fig. 19) by the presence of a clear, short and narrow scape (Sp), almost absent in the other species, by the shorter fertilization ducts (Fd) with fewer coils, and the distally wider copulatory ducts (Cd) (Fig. 58F vs. Fig. 62F). A further diagnostic character is the uniformly dark color of the opisthosoma lacking any pattern ( Fig. 58A-C vs. Fig. 62A-C).
Description. Habitus as in Fig. 58A-C. Carapace pale yellow, with dense, dark marks. Cervical groove, fovea and radial furrows distinct. Thoracic area dark at its margins. Mouthparts pale yellow, faintly pigmented. Sternum with a very sharp posterior corner, with a reticulated and finely pigmented surface. Legs and female palps pale, lacking spines. Opisthosoma black, densely covered with setae. Epigyne (Fig. 58D-F): light-colored (Fig. 58D). Scape short and narrow, rectangular and slightly protruding beyond the epigynal posterior margin (Fig. 58D-E). Spermathecae weakly sclerotized, ovoid, separated by about two diameters (Fig. 58E). Fertilization ducts thin and long, forming a small and a large loop before reaching the spermathecae (Fig. 58F). Copulatory ducts thick and dark, firstly bent inward and then outward, mesially swollen and distally narrower (Fig. 58E-F Etymology. The specific name is derived from the type locality; adjective. Diagnosis. The new species is closely related to Nesticella connectens (see Figs 50A-D, 51A-G), N. nepalensis (see Figs 55A-D, 56A-G) and N. potala sp. n. (see Fig. 57A-F). Males can be recognized by the curved process II of the ventral apophysis (Va-II), the stockier distal process II (Dp-II) and the different shape of the terminal apophysis (Ta) (Fig. 59A-B, D vs. Fig. 50A-B, D vs. Fig. 55A-B, D). Females can be distinguished from those of N. connectens by the more coiled fertilization ducts (Fd) and the shorter copulatory ducts (Cd) (Fig. 60G vs. Fig. 51G); from those of N. nepalensis by the longer scape (Sp), the wider spermathecae (S) and the more twisted fertilization ducts (Fd) (Fig. 60E-G vs. Fig. 56E-G); and finally from those of N. potala sp. n. by the almost round spermathecae (S), the 90° angle of the copulatory ducts (Cd) (approx. 45° in N. potala sp. n.) and by the presence of a clear abdominal pattern which is absent in the other species (Fig. 60E-G vs. Fig. 57D-F).
Description. Habitus as in Fig. 60A-D. Carapace yellowish in males, darker in females. Cervical groove distinct, fovea deep. Thoracic area with weak radial furrows. Mouthparts pale yellow. Sternum greyish. Legs and female palps yellowish, distally darkish in metatarsi and tarsi. Opisthosoma covered with long setae, yellowish with paired dark marks, partially merged each other on the dorsal area and bordering a cross-like light mark.
Habitat. Forest leaf litter. Distribution. Known only from the type locality (Fig. 83).   Etymology. The specific name is derived from Van Lang, an ancient name for Vietnam; noun in apposition.
Diagnosis. 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. 61E-F). N. vanlang sp. n. can be distinguished from N. sogi (see Lehtinen and Saaristo1980: 57, fig. 21) by the smaller size of the spermathecae (S) and the longer copulatory ducts (Cd) (Fig. 61F vs. fig. 21).
Description. Habitus as in Fig. 61A-C. Carapace greyish and faintly pigmented. Cephalic area with several long setae along the midline and the thoracic groove. Cervical groove and fovea distinct. Thoracic area pigmented at margins. Mouthparts pale yellow. Labium very wide. Sternum smooth and greyish. Legs and female palps yellowish, metatarsi and tarsi distally darker. Opisthosoma pale greyish, darker ventrally. Epigyne (Fig. 61D-F): strongly clerotized, reddish-brown (Fig. 61E). Scape rectangular, very short and barely visible, (Fig. 61E-F). Copulatory openings near the posteromargin of the epigyne, separated from each other by more than a half of the spermathecal diameter ( Fig. 61E-F). Spermathecae almost round, close to each other and separated by less than a half of their diameter (Fig. 61F). Fertilization ducts thick, coiled into a single loop before reaching the spermathecae (Fig. 61E-F). Copulatory ducts thick and short (Fig. 61F).

Habitat. Forest leaf litter.
Distribution. Known only from the type locality (Fig. 83).
Habitat. Forest leaf litter. Distribution. China (Yunnan) (Fig. 83) Etymology. The new species is named after Dr. Zhiyuan Yao who extensively collected and studied spiders from Southeast Asia; noun (name) in genitive case.
Diagnosis. 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. 63A-B, D). Females can be easily recognized by the strongly reduced, almost absent, scape (Sp), by the piriform spermathecae (S) and by the short copulatory ducts (Cd) (Fig. 64E-G) which are not shared with any other species of the group.
Description. Habitus as in Fig. 64A-D. Carapace yellowish in males, darker in females; dark area present before the fovea. Cervical groove and fovea distinct. Thoracic area dark, mouthparts yellow. Legs and female palps yellowish, distally darker in metatarsi and tarsi, faintly dark in the other segments. Opisthosoma dark with a dorsal vertical pale stripe and a posterior irregular spot, covered with long setae.

Nesticella phami-group
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. Diagnosis. Males of the new species can be distinguished from those of Nesticella sumatrana sp. n. (Fig. 67A-G) by the strongly sclerotized terminal apophysis (Ta), the clearly serrated tegular apophysis (Tg) (Fig. 65A, C vs. Fig. 67A, C), the longer, and wider distal process of the paracymbium (Dp) (Fig. 65A-B, D vs. Fig. 67A-B, D) and the reduced ventral apophysis (Va) (Fig. 65A vs. Fig. 67A). Females can be easily separated from those of the other Nesticella species by the very short and wide scape (Sp), the short fertilization and copulatory ducts (Fd and Cd) and the wide, ovoid spermathecae (S).     Fig. 66A-D. Carapace uniformly yellow and faintly pigmented at center and at margins. Cervical groove faint, fovea shallow. Mouthparts brown-yellowish. Sternum with sparse long setae and shorter setae along margins. Legs and female palps yellowish, distally brownish in metatarsi and tarsi. Opisthosoma pale in males, darker in females, with paired black marks on the dorsal part, partially fused each other in the posterior side.

Nesticella quelpartensis-group
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 Paik & Namkung, in Paik et al. 1969: 812, figs 37-42) by the shorter ventral apophysis of the  Description. Habitus as Fig. 69A-D. Carapace pale yellow in males, darker in females, faint dark at margins and near the center. Cervical groove and fovea distinct. Mouthparts brown-yellowish. Sternum yellow, with sparse long setae. Legs and female palps yellowish, distally darker in metatarsi and tarsi. Opisthosoma light yellow with paired dark marks, partially merged with each other on the posterior, forming a light, cross-shaped mark on the background. Pattern fainter in males.

Nesticella quelpartensis
Description. See Paik et al. (1969).  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.
Description. 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.
Diagnosis. 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 Liu and Li 2013a: figs 1-2, 7-8). Females of this species can be distinguished by the wider, piriform spermathecae rather than small and almost round spermathecae, by the U-shaped scape (Sp) and by the smaller vulval pockets (Vp) (respectively slimmer, triangular or U-shaped but with a larger proximal part and wider in the other species) (see Liu and Li 2013a: figs 4-5, 12-14).
Description. See Liu and Li (2013a (Fig. 76D-G). Males of this species can be distinguished from those of P. spinosus sp. n. by the longer dorsal apophysis (Da) (Fig.  70A-B, D vs. Fig. 74A-B, D), the compact distal process (Dp) lacking any serrated margin (Fig. 70D vs. Fig. 74D), the absence of a clear additional tegular apophysis (Tg-II) (Fig. 70A vs. Fig. 74A), the sharper and longer terminal apophysis (Ta) (Fig.  70A vs. Fig. 74A) and the wider conductor with two clear processes (Cp-I and Cp-II) (Fig. 70C-D vs. Fig. 74C-D). Females can be distinguished from those of P. spinosus sp. n. by the smaller, triangular scape (Sp), the ducts (Fd and Cd) closer to each  other and the darker coloration (Fig. 71E, H vs. Fig. 75E, H); from those of P. wumengensis sp. n. by the narrower epigynal plate (EP) (Fig. 71E-F vs. Fig. 76D-E) and the shorter and more triangular scape (Sp) (Fig. 71G-H vs. Fig. 76F-G). The narrow, triangular shape of the scape (Sp) with a sharp tip and the general configuration of the ducts (Fd and Cd), strongly coiled, allow an easy separation of P. dafangensis sp. n. from all other species in this genus.
Description. Habitus as in Fig. 71A-D. Carapace pale yellow, with dark marks at margins and around cephalic area. Eyes ALE>PLE>PME>AME. Cervical groove and fovea distinct. Thoracic area with faint radial furrows. Mouthparts pale yellow, darker in females. Sternum yellowish, lightly pigmented. Legs and female palps yellowish, distally darker in metatarsi and tarsi. Opisthosoma very dark, especially in males, with lighter horizontal strips.
Male (   Etymology. 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. Diagnosis. The new species is closely related to P. dafangensis sp. n. (Figs 70A-D, 71E-H) and P. wumengensis sp. n. (Fig. 76A-G). Males can be easily separated from those of the former species by the spiked distal process of the paracymbium (Dp), the shorter dorsal apophysis (Da) (Fig. 74A-B, D vs. Fig. 70A-B, D), the presence of a secondary tegular apophysis (Tg-II) (Fig. 74A vs. Fig. 70A) and the stouter, terminal apophysis (Ta) (Fig. 74A vs. Fig. 70A). Females can be distinguished from those of P. dafangensis sp. n. by the wider arrow-like scape (Sp), the ducts (Fd and Cd) closer to each other and the lighter coloration ( Fig. 75A-H vs. Fig. 71A-H). They can be recognized from those of P. wumengensis sp. n. by the wider and more triangular scape (Sp) with a more pointed tip (Fig. 75E-H vs. Fig.  76D-G). The spiny distal process of the paracymbium (Dp) and the triangular, pointed epiginal scape (Sp) allow easy separation from all the other species of Pseudonesticus.
Male ( Fig. 71E-H) and P. spinosus sp. n. (see Fig. 75E-H). It can be distinguished from these two species by the longer and narrower scape (Sp), the wider distance between the spermathecae (S) and the wider epigynal plate (Ep) (Fig. 76E-G vs. Figs 71F-H, 75F-H). The same combination of characters allows to separate P. wumengensis sp. n. from all the other species of Pseudonesticus.
Female (    Etymology. The specific name is derived from the type locality; adjective. Diagnosis. The new species is closely related to P. miao sp. n. (Figs 72A-D, 73F-H) and P. clavatus (see Liu and Li 2013a: 790, figs 1-14). Males can be distinguished from those of the former species by the shorter and undulate terminal apophysis (Ta), the longer process I of the conductor (Cp-I) and the pointed distal process of the paracymbium (Dp) (Fig. 77A-B, 77D vs. Fig. 72A-B, D). Females can be distinguished from those of P. miao sp. n. by the shorter and wider scape (Sp) (Fig. 78E-G). Males of P. ziyunensis sp. n. can be separated from those of P. clavatus by the shorter terminal apophysis (Ta), the longer distal process of the paracymbium (Dp), the shorter process I of the conductor (Cp-I), the lack of a ventral apophysis (Va); females are distinguished by the shorter and wider scape (Sp), the smaller spermathecae (S), the coiled ducts (missing any coil in the other species) and the wider and cystiform vulval pockets (Vp) (Figs 77A, 78E-G vs. figs 1, 12-14). The general shape of the paracymbium (P), the long terminal apophysis (Ta) and the general shape of the scape (Sp) allow an easy separation from all other species of Pseudonesticus.
Male ( Etymology. 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. Diagnosis. 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 welldeveloped and strongly protruding in the latter genus.
Male palp (Fig. 79A-D): tibia elongate ( Fig. 79A-B). Paracymbium well-developed, sickle-like, with a single ventral apophysis usually short and squared; a short, flat dorsal apophysis and a sclerotized distal process with two ramifications (Fig. 79A-B, D). Terminal apophysis triangular or rectangular, elongate and protruding prolaterally. Tegular apophysis reduced (Fig. 79C). Conductor wide and convoluted, with three distinct processes, two elongate and one flat and laminar (Fig. 79A). Embolus filamentous, starting from the posterior side of the bulb and reaching the apex of the conductor with a half loop (Fig. 79A, C). Epigyne (Fig. 80C-D): broad, with a very short scape (Fig. 80C). Well sclerotized ducts partially visible through the tegument. Copulatory openings wide, located at the lateral side of the scape (Fig. 80C). Spermathecae small and almost round (Fig. 80D). Fertilization and copulatory ducts short and slightly convoluted (Fig. 80D). Vulval pockets well-developed, with a wide, balloon-like shape, located above the spermathecae (Fig. 80D).
Distribution. China (Guangxi, Guizhou, Hunan), Japan (Honshu Island, Shizuoka Prefecture).     Remarks. 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.   Wunderlich et Song, 1995 11 N. zhiyuani sp. n