On the clubionid spiders (Araneae, Clubionidae) from Xishuangbanna, China, with descriptions of two new genera and seven new species

Abstract Clubionid spiders from Xishuangbanna, Yunnan Province, China are studied. A total of seven genera and 13 species have been found, including two new genera with one new species each, i.e., Ramosatidia Yu & Li, gen. nov., with R.situ Yu & Li, sp. nov. (♂♀) as the type species and Sinostidia Yu & Li, gen. nov., with S.shuangjiao Yu & Li, sp. nov. (♂♀) as the type species. Five additional new species are Sinostidiadujiao Yu & Li, sp. nov. (♂♀), Matidiaxieqian Yu & Li, sp. nov. (♂♀), Nusatidiachangao Yu & Li, sp. nov. (♂♀), N.mianju Yu & Li, sp. nov. (♀), and N.subjavana Yu & Li, sp. nov. (♀). The following genera and species are reported from China for the first time: Malamatidia Deeleman-Reinhold, 2001, Pteroneta Deeleman-Reinhold, 2001, Malamatidiazu Jäger & Dankittipakul, 2010, Nusatidiaaeria (Simon, 1897), N.camouflata Deeleman-Reinhold, 2001, Porrhoclubionapteronetoides (Deeleman-Reinhold, 2001), and Pteronetaultramarina (Ono, 1989). Malamatidiachristae Jäger & Dankittipakul, 2010 syn. nov. is a junior synonym of Malamatidiazu. Nusatidiarama Deeleman-Reinhold, 2001 syn. nov. is synonymised with N.aeria (Simon, 1897).


Introduction
The Clubionidae Wagner, 1887 is a relatively large family with 656 valid species distributed worldwide (WSC 2021), and 517 of these are placed in the genus Clubiona Latreille, 1804(WSC 2021. Before the current study, 178 species in five genera (Li 2020) were known from China, with 172 species assigned to Clubiona. Of these, 51 Clubiona species have been recorded from Xishuangbanna, with 30 described as new species in the last three years (Yu and Li 2019a, b;Zhang et al. 2021). Herein, we discuss 13 species in seven genera of Clubionidae found in Xishuangbanna, including two genera with seven species new to science, and two genera with five species new to China.

Materials and methods
Specimens were primarily collected by canopy fogging, while a few were obtained by beating vegetation and pitfall trapping. All type specimens are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS, curator Jun Chen) in Beijing, China.
Specimens were examined using a LEICA M205C and an Olympus SZX7 stereomicroscope. Further details were studied under a CX41 compound microscope. Male and female genitalia were examined and illustrated after dissection. Left male palps are illustrated, unless otherwise indicated, and photos of the right palps are flipped horizontally in figures to allow ease of comparison with other species. Epigynes were removed and cleared in lactic acid or warm 10% potassium hydroxide (KOH) solution before illustration. Vulvae were imaged after being embedded in Arabic gum (Yuanye Biotechnology Co., Ltd). Images were captured with a Canon EOS 70D digital camera (20.2 megapixels) mounted on an Olympus CX41 compound microscope and assembled using Helicon Focus 6.80 image stacking software. All measurements were obtained using an Olympus SZX7 stereomicroscope and are given in millimetres. Eye diameters are measured at the widest points. The total body length does not include the chelicerae or spinnerets. Leg lengths are given as total length (femur, patella+tibia, metatarsus, tarsus).
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). Abbreviations used in the text and figures are as follows: Most of the terminology used in the text and figure legends follows Yu and Li (2019a, b), and some follows Deeleman-Reinhold (2001) and Zhang et al. (2018).
Sequences were trimmed to 656 bp. All sequences were confirmed using BLAST and are deposited in GenBank. The codes and GenBank accession numbers of voucher specimen are provided as follows: Malamatidia zu: YHCLU0119, ♂, GenBank MZ508477; YHCLU0120, ♀, GenBank MZ508476. Matidia spatulata Chen & Huang, 2006 Embolus shorter than half of tegulum, its free part slightly curved, tip directed antero-retrolaterally; tegular apophysis present; sperm duct bulky and twisted; RTA concave with two branches (Fig. 7A-E); epigyne ventrally with a circular or square posterior plate; copulatory ducts short and not convoluted; spermathecae large and elongate, bursae reduced or absent ( Fig. 8A-D); metatarsi I and II with a pair of dorso-basal spines .............. N. aeria and N. luzonica -Embolus equal to or longer than tegulum length, typically oriented clockwise along the margin of the tegulum, or draped around the tegulum; tegular apophysis reduced or absent, or represented by a membranous prolongation; RTA simple, unbranched; epigyne without posterior plate; copulatory ducts relatively long, usually convoluted; both spermathecae and bursae present, both subglobular ( (cf. Figs 2E,G and 4E,G,6E,G) and femur I is shorter than femur II. It differs from Nusatidia by the sternum without a rectangular extension (cf. Figs 2H and 10B). Malamatidia species also can be recognised by the copulatory organs: RTA with a sub-rectangular tip (Fig. 1B), filiform embolus curving clockwise and following along the tegular distal margin (Fig. 1C, D), sperm duct an S-shaped loop and sometimes abruptly narrowed (Fig. 1D, E); epigynal plate has a central depression with a longitudinal rim or septum centrally ( Fig. 2A, B). All characters of Malamatidia, Nusatidia, and Matidia are according to Deeleman-Reinhold (2001) and Jäger and Dankittipakul (2010).
Description. See Deeleman-Reinhold (2001). Comments. Based on morphological characters, the genus is probably closely related to Matidia and Nusatidia. However, the monophyly and the exact placement of Malamatidia within Clubionidae remains to be tested. Diagnosis. Males can be characterised by the sickle-shaped embolus, its tip terminated antero-mesally (Fig. 1C, D) (vs. embolus filiform or flagelliform, terminated postero-retrolaterally in other Malamatidia spp.). The female of M. zu is easily differentiated from all other congeners by the oval bursae (Fig. 2C, D) (vs. globular bursae).

Malamatidia zu
Description. See Jäger and Dankittipakul (2010). Male palp as in Fig. 1A-E, epigyne as in Fig Comments. Both sexes were known for all Malamatidia species except M. christae and M. zu. These two species were described based on holotypes from Laos. The former was collected from Luang Nam Tha Province, while the latter was from Luang Prabang Province. Jäger and Dankittipakul (2010)  Diagnosis. Species in this genus differ from all other clubionids by the following: the pars cephalica is 2× narrower than the pars thoracica (Figs 4E, G, 6E-G) (vs. wider), leg I is longest (vs. not longest), and there is a dark ventral abdominal spot in males ( Fig. 4F) (vs. absent). Matidia resembles Malamatidia, Nusatidia, and Ramosatidia gen. nov. by the slender, greenish body but is consistently separable by the shape of the copulatory organs: the male palp has a ribbon-shaped embolus (Figs 3A, C-F, 5A, C-F) (vs. embolus not ribbon-shaped), the epigyne has one or two depressions (or an atrium) and no septum (Figs 4A,B,6A,B) (vs. depression lacking in Nusatidia and Ramosatidia gen. nov., or present but with a septum in Malamatidia).  Comments. Based on the two newly discovered species, the description should be extended from Deeleman-Reinhold (2001): the epigyne of M. spatulata Chen & Huang, 2006 with 2 depressions, copulatory ducts relatively long (longer than epigyne) (Fig. 4A-D) (vs. one central atrium and short ducts (shorter than epigyne) in all other congeners (e.g., M. xieqian sp. nov.; Fig. 6A-D)); an additional exceptional feature in M. xieqian sp. nov. is that the cheliceral promarginal teeth are farther from the fang base than the retromarginal ones (vs. promarginal teeth nearer the fang base in all other known Matidia species). Both M. spatulata and M. xieqian sp. nov. have anteriorly located bursae (vs. bursae located laterally or posteriorly).
Note. Deeleman-Reinhold (2001) considered Matidia putatively polyphyletic. In the same work, she established three new, similar closely related genera (Nusatidia, Pristidia, and Malamatidia) to accommodate 12 new species from SE Asia, and placed them in the subfamily Clubioninae. She also transferred four Matidia species to Nusatidia. The similar somatic characters and sympatric distribution strongly suggest close relationships between the four genera. However, the phylogenetic relationships remain unresolved (Versteirt et al. 2010).     Etymology. The specific name is derived from the Chinese pinyin xiè qián, which means crab claw, referring to the concave retrolateral tibial apophysis with two branches resembling a crab claw; noun in apposition.
Distribution. Known only from the type locality.

Genus Nusatidia Deeleman-Reinhold, 2001
Nusatidia Deeleman-Reinhold, 2001: 166. Comments. The somatic characters of Nusatidia species strongly suggest a close relationship with Matidia. However, both genera are possibly paraphyletic (Versteirt et al. 2010). According to WSC (2021), only two described Nusatidia species were known from males: N. luzonica (Simon, 1897) and N. manipisea (Barrion & Litsinger, 1995), both from Luzon Island in the Philippines. We cannot rule out the possibility that these two species are conspecific to C. mianju sp. nov. and C. subjavana sp. nov. Comments. Nusatidia aeria was originally described in Matidia based on the holotype female from Jolo Island, Philippines. Deeleman-Reinhold (2001) examined the holotype and transferred the species to Nusatidia. In the same work, she described N. rama based on the holotype male from Sumatra but suggested that these two species could be conspecific. Recently, new material has been collected from Xishuangbanna containing both sexes. The males were identified as N. rama while the females were identified as N. aeria. On the basis of the morphological characters ( Fig. 8E-H) and DNA barcoding, we matched the females and males. Therefore, the two species are synonymised, and priority is given to N. aeria.

Nusatidia aeria
Distribution. Prior to our study, this species was known from Borneo and Indonesia (Sumatra) only. Our collection in southwest China (Yunnan Province, new record) extends the known range of this species ~ 2700 km to the northwest.   Distribution. Prior to our study, this species was known from Thailand (Kanchanaburi Province) only. Our collection in southwest China (Yunnan Province, new record) extends the known range of this species ~ 870 km to the northwest. Etymology. The specific name is derived from the Chinese pinyin cháng áo, meaning long chelicerae, referring to the enlarged chelicerae of the male, which are approximately as long as the carapace; noun in apposition.
Diagnosis. Males of N. changao sp. nov. resemble those of N. borneensis Deeleman- Reinhold, 2001 andN. snazelli Deeleman-Reinhold, 2001 in having a similar embolus draped around the tegulum but differ by the tibial apophysis, which has a flange with jagged teeth like those on a saw (  be easily recognised by the enlarged chelicerae, ~ as long as the carapace (Fig. 12E) (vs. chelicerae unmodified, < 1/2 of carapace length).
Distribution. Known only from the type locality. Etymology. The specific name is derived from the Chinese pinyin miàn jù, for mask, referring to the conspicuousness of the spermathecae and copulatory ducts through the epigynal plate, the general appearance of a mask; noun in apposition.
Male. Unknown. Distribution. Known only from the type locality. Other material examined. 1♀ (YHCLU0132), same data as holotype.

Nusatidia subjavana
Etymology. The specific name is a Latin adjective referring to the species' similarity to N. javana (Simon, 1897), a combination of the preposition sub (near) and the epithet of that species.
Male. Unknown. Distribution. Known only from the type locality.

Comments.
Porrhoclubiona is small Clubionidae with a relatively wide body and can be easily separated from Malamatidia, Matidia, and Nusatidia. Porrhoclubiona currently comprises two species groups, the pteronetoides group and the genevensis group. The pteronetoides group was established by Deeleman-Reinhold (2001) based on two species endemic to the Oriental realm. The genevensis group was formally named by Mikhailov (1992), though in fact it was first recognised by Lohmander (1944) as a subgenus of Microclubiona (currently considered junior synonyms of Clubiona), and then refined by Bosmans et al. (2017) with eight species from West Palaearctic region. The two groups share almost all of the generic characters listed by Marusik and Omelko (2018) (see diagnosis above); however, pteronetoides group can be differed from the genevensis group by have a dorsal abdominal scutum (Fig. 16E, F) (vs. absent), lack modified setae on the cymbium (Fig. 15B) (vs. present) in males, and the spermathecae are elongate in females (Fig. 16C, D)  Diagnosis. Both sexes of P. pteronetoides are similar to those of P. viridula in having similar palps and epigynes but can be differentiated by the elongate-oval bulb in ventral view (Fig. 15B) (vs. pyriform or triangular) and by the oval spermathecae that are ~ 2× longer than wide (Fig. 16C, D) (vs. claviform spermathecae that are ~ 3-4× longer than wide).
Description. See Deeleman-Reinhold (2001). Male palp as in Fig. 15A-E, epigyne as in Fig. 16A-D, habitus as in Fig. 16E-H. Comments. There is almost no difference between males from Xishuangbanna ( Fig. 15A-E) and the holotype from Thailand (Deeleman-Reinhold 2001: 97, figs 5-7). However, some intraspecific variation is exhibited by females from the two localities, related to distance between the two copulatory openings. The copulatory openings are separated by ~ 1/4 of their diameter in material from Xishuangbanna (    E, F, equal for G, H).

Pteroneta ultramarina (Ono, 1989) Figs 17, 18
Clubiona ultramarina Ono, 1989: 156, figs 1-7 (♂♀).  (Versteirt et al. 2008: 312, fig. 7a, b) in having a similarly shaped retrolateral tibial apophysis and claw-like embolus but differ by the conductor with a straight tip (Fig. 17A-E) (vs. semi-circular tip) and the dorsal surface of the chelicerae with relatively few, short setae (< 10) (Fig. 18E) (vs. 18 short setae). Females of P. ultramarina are similar to those of P. tertia Deeleman-Reinhold, 2001. The epigyne of these two species is very similar and almost indistinguishable, but the species differ in the number and arrangement of cheliceral teeth (three promarginal and two retromarginal in P. ultramarina vs. six teeth on both margins in P. tertia) and by the patterns on the body (the carapace and sternum are marked with lazulite-coloured blue spots in P. ultramarina (Fig. 18G, H) (vs. blue spots lacking in P. tertia).
Description. See Ono (1989). Male palp as in Fig. 17A-E, epigyne as in Fig.  18A-D, habitus as in Fig. 18E-H. Distribution. Japan (Ryukyu Is.), China (Yunnan Province, new record). The new record presented here extends the known range of this species by ~ 2700 km from the type locality (Ryukyu Is.) to the southwest (Xishuangbanna).   E, F, equal for G, H).

Ramosatidia Yu & Li, gen. nov.
http://zoobank.org/0DA81522-8B4E-41B4-8325-03D62825768F Type species. Ramosatidia situ Yu & Li, sp. nov. Etymology. The generic name is a combination of the Latin adjective ramosus, which means ramose, or branching, referring to the apophyses of the palpal tibia, in conjunction with atidia, alluding to the green colour and slender body, similar to Matidia. The gender is feminine.
Diagnosis. Ramosatidia gen. nov. resembles the other genera exclusively distributed in SE Asia (Pristidia, Nusatidia, and Matidia) by the similar habitus (green, elongate, long-legged), but it is consistently separable by somatic characters and the copulatory organs. This new genus is characterised by the promargin with only one tooth in the male and the retromargin without tooth in the female and by the bottle-green body in in ethanol (vs. living spiders are pale green, but specimens are pale yellow, white or brownish in ethanol in almost all other genera). Ramosatidia gen. nov. can be distinguished from Pristidia by the relatively small eyes, with the PME > their diameter apart (Fig. 20E, G) (vs. PME barely > their diameter apart), from Nusatidia by the sternum lacking a rectangular extension beyond coxae I (cf. Fig. 20H and 10B), and from Matidia by the higher ocular region/carapace width ratio (approximately 2/3 vs. 1/2 in Matidia) (cf. Figs 20E,G and 4E,G), and femur I shorter than femur II (vs femur I longest). Ramosatidia gen. nov. species also can be recognised by the following characters of the copulatory organs: male palpal tibia with four apophyses (vs. maximum of three apophyses) (Fig. 19A-E); epigynal plate lacking atrium, depression, and septum (vs. with atrium or depression in Matidia, with septum in Malamatidia), fertilisation ducts relatively large, ~ 1/2 of spermathecae length (Fig. 20A-D) (vs. shorter than 1/2 of spermathecae length). All the provided characters of Pristidia, Nusatidia, and Matidia are according to Deeleman-Reinhold (2001) and recent clubionid papers, such as Yu et al. (2012Yu et al. ( , 2017. Description. Same as for the type species. Composition. Type species only. Distribution. China (Yunnan). Other material examined. 1♀ (YHCLU0134), same data as holotype.

Ramosatidia situ
Etymology. The specific name is derived from the Chinese pinyin sìtū, which means four apophyses, referring to four tibial apophyses; noun in apposition.
Distribution. Known only from the type locality. Diagnosis. This genus can be easily confused with Pristidia due to a similar appearance. Sinostidia gen. nov. and Pristidia share a similar cephalic region/carapace width ratio, relatively large eyes (PME barely > their diameter apart), tibial spination, and pale brownish body, but they can be separated by Sinostidia gen. nov. having promarginal teeth closer to the fang base than the retromarginal ones. The copulatory organs of Sinostidia gen. nov. resemble those of Pristidia in having a similar bulb with a sharply pointed embolus arising dorsally, hidden by the tegulum and by having a similar epigynal plate, but differ by: (1)  Description. Medium-sized, with body length of males 4.5-5.0, females 4.79-5.6. Body yellow-white, legs uniformly coloured as carapace. Carapace: elongate-oval in dorsal view, pars cephalica slightly elevated above thorax, pars thoracica distinctly wider than pars cephalica; integument smooth, with sparse, erect, thin, dark bristles on pars cephalica (bristles detach easily in ethanol); yellow or pale orange, slightly darker in ocular region, without distinct pattern; fovea longitudinal, reddish. Clypeus height distinctly less than diameter of AME. Chelicerae robust, brownish red, fang furrow with three promarginal and two retromarginal teeth. Sternum yellowish, anteriorly straight, anterior and lateral margins with brown extensions fitted into intercoxal concavities; posterior region strongly protruded between coxae IV. Eyes: arranged in a compact group; AER slightly recurved in dorsal view, procurved in anterior view, AME very slightly smaller than ALE, or equal in diameter, AME closer to ALE than to each other; PER recurved in dorsal view, PME distinctly larger than PLE, PME separated by one diameter. Legs: formula usually IV, I, II, III; all tarsi scopulate; anterior metatarsi with a pair of basal ventral spines; tibiae I and II with two pairs of strong ventral spines; tibiae and metatarsi of posterior legs with more spines than anterior legs, but spination varies among different individuals. Abdomen: lanceolate, tapered posteriorly, uniformly white, dorsum with numerous indistinct patches, or anteriorly with a longitudinal, grey heart mark; venter, sides white, without distinct markings. Spinnerets: anterior lateral spinnerets short and coniform; posterior lateral spinnerets cylindrical, relatively long; anterior median spinnerets small, sandwiched between anterior lateral spinnerets and posterior lateral spinnerets. Male palp: Femur and patella unmodified. Tibia short, no longer than 1/2 of cymbium length, with two apophyses: ventral apophysis stout, with blunt tip, typically thumb-like; retrolateral apophysis weak, shape variable, tip relatively pointed. Cymbium unmodified, ~ 1.8× longer than wide, with dense dorsal setae. Bulb elongateoval, embolic area located distally on tegulum. Tegular apophysis large, longer than 1/2 of tegulum width, arising at ~ 1 o'clock position, gradually tapered toward apex, pointed prolatero-distally, covering embolic base. Embolus with bulky base and sharp tip, curved behind tegular apophysis, tip extended to apex of cymbium. Conductor small, situated retrolaterally on tegulum.
Comments. The large PME and the presence of a claw-shaped embolus located behind the tegulum indicate that the new genus is likely closely related to Pristidia. Somatic characters are either poorly delineated or variable, making the differentiation of Sinostidia gen. nov. and Pristidia difficult. However, the two new species share a distinct set of genitalic characters and can be easily separated from Pristidia, thus, we established a new genus to accommodate them.  Other material examined. 1♂ (YHCLU0132), same data as the holotype. Etymology. The specific name is derived from the Chinese pinyin dú jiǎo, which means one-horned, referring to the unbranched tegular apophysis; noun in apposition.
Diagnosis. Males of the new species are similar to those of S. shuangjiao sp. nov. but can be distinguished by the unbranched tegular apophysis, wide and conical embolar tip, and by the thick and partly membranous conductor ( Fig. 23A-E) (vs. tegular apophysis bifurcate, embolar tip filamentous, conductor thin and entirely membranous as in Fig. 21A-E). Females of S. dujiao sp. nov. differ from those of S. shuangjiao sp. nov. by the reniform bursae larger than spermathecae (Fig. 24C, D) (vs. globular bursae smaller than the spermathecae as in Fig. 22C, D), the CO situated anteriorly and the CD located on central portion of epigyne (vs. both situated laterally in S. shuangjiao sp. nov. ).
Distribution. Known only from the type locality.