Research Article |
Corresponding author: Shuqiang Li ( lisq@ioz.ac.cn ) Academic editor: Yuri Marusik
© 2015 Shuqiang Li, Sergei Zonstein.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Li S, Zonstein S (2015) Eight new species of the spider genera Raveniola and Sinopesa from China and Vietnam (Araneae, Nemesiidae). ZooKeys 519: 1-32. https://doi.org/10.3897/zookeys.519.8784
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Eight new species, seven Raveniola Zonstein, 1987 and one Sinopesa Raven & Schwendinger, 1995 from China and Vietnam are described: Raveniola alpina sp. n., R. bellula sp. n., R. chayi sp. n., R. gracilis sp. n., R. rugosa sp. n., R. spirula sp. n. and R. yajiangensis sp. n. and Sinopesa ninhbinhensis sp. n. Keys to all East-Asian congeners, diagnoses of the new species, and new distribution data of R. montana Zonstein & Marusik, 2012, with a first record for Sichuan, China, are provided.
Taxonomy, mygalomorph spiders, new records, South-East Asia
The Nemesiidae are known to include 45 genera and 382 species (
While sorting and identifying nemesiid material in the Institute of Zoology, Chinese Academy of Sciences, eight new species belonging to Raveniola and Sinopesa were found; they are described here. The systematic position of both genera was recently considered by
To permit reliable identification and stimulate further studies in this area, all new descriptions are illustrated and keys to the studied genera and species are added.
Specimens were examined and measured with a LEICA M205 C stereomicroscope and details were studied with an Olympus BX51 compound microscope. Illustrations were made using a camera lucida attached to the Olympus BX51 microscope and inked with an ink jet plotter. Male palps and female genitalia were examined and illustrated after they were dissected from the spiders. Vulvae were treated in a warm solution of 10% potassium hydroxide (KOH). The left palp and left legs I and II of male spiders were illustrated, unless otherwise indicated. Specimens were preserved in a 75% ethanol solution. Photos were taken with an Olympus C7070 wide zoom digital camera (7.1 megapixels) mounted on an Olympus SZX12 stereomicroscope. The images were assembled using Helicon image stacking software. All measurements are given in millimetres unless otherwise noted. Leg measurements are given as: total length (femur + patella + tibia + metatarsus + tarsus). Leg segments were measured on the dorsal side.
The following abbreviations are used: AL – abdomen length; ALE – anterior lateral eye(s); AME – anterior median eye(s); AW – abdomen width; CL – carapace length; CW – carapace width; PLE – posterior lateral eye(s); PLS – posterior lateral spinneret(s); PME – posterior median eye(s); PMS – posterior median spinneret(s); TL – total length (including chelicerae, but not spinnerets).
All specimens used in this study are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing, China. The only exception is the female paratype of R. chayi sp. n., kept at the Senckenberg Museum, Frankfurt am Main, Germany (SMF).
Only two genera of the family occur with some degree of certainty in eastern Asia. Judging from the original description, Nemesia sinensis Pocock, 1901 probably belongs to the Cyrtaucheniidae (see
The distribution of Atmetochilus is given considering data provided by
1 | Thoracic fovea short, U-shaped; posterior sternal sigilla distinctly larger and farther from sternal margin than other sigilla, in many cases subcentral or confluent; paired tarsal claws either with two more or less distinctly juxtaposed teeth rows (females) or with one S-shaped row (males); PMS well developed; male tibia I with coupling spur and megaspines located proventrally | 2 |
– | Thoracic fovea short, straight or pit-like; posterior sternal sigilla submarginal; paired tarsal claws with two similar teeth rows on promargin and retromargin in males and females; PMS small to absent; male tibia I with two enlarged retroventral distal spines and without coupling spur | 4 |
2 | Posterior sternal sigilla submarginal to subcentral but not confluent (Raven 1985: fig. 53; |
3 |
– | Posterior sternal sigilla confluent; (Raven 1985: fig. 59) | Atmetochilus (India, Indonesia, Myanmar and Thailand) |
3 | Metatarsal preening combs absent on legs III and IV; female tarsus IV with scopula | Damarchilus (Eastern India) |
– | Metatarsal preening combs present on legs III and IV; female tarsus IV without scopula | Damarchus (Eastern India, Indonesia, Malaysia, Myanmar, Singapore and Thailand) |
3 | Carapace hirsute and with finely granular texture; hairs on legs I–IV long and non-uniform; tarsal scopula more or less dense and long; male intercheliceral tumescence reduced if present | Raveniola (south Palearctic, from Turkey to China) |
– | Carapace with only a few bristles; hairs on legs I–IV more or less uniformly short; tarsal scopula thin and short; male intercheliceral tumescence well developed | Sinopesa (South-eastern China, Ryukyu Isles, Thailand and Vietnam) |
Brachythele virgata Simon, 1891, from Central Asia, by the original designation.
Raveniola, similar to Sinopesa Raven & Schwendinger, 1995, has two enlarged retroventral distal spines on tibia I in males and divided receptacles in females, as well as the absence of a serrula and metatarsal preening combs. The leg scopula in Raveniola is more developed than it is in Sinopesa. By contrast, the male intercheliceral tumescence in Raveniola is lost or vestigial, whereas in Sinopesa, it is well-developed. Like Sinopesa, members of Raveniola have more or less reduced PMS, which are completely lost in some species. The apical segment of the PLS in Raveniola is usually shorter than that in Sinopesa.
Raveniola currently comprises 29 species, including the new species described here; 14 of them occur in China: R. alpina sp. n., R. bellula sp. n., R. chayi sp. n., R. gracilis sp. n., R. guangxi (Raven & Schwendinger, 1995), R. hebeinica Zhu, Zhang & Zhang, 1999, R. montana Zonstein & Marusik, 2012, R. rugosa sp. n., R. shangrila Zonstein & Marusik, 2012, R. songi Zonstein & Marusik, 2012, R. spirula sp. n., R. xizangensis (Hu & Li, 1987), R. yajiangensis sp. n. and R. yunnanensis Zonstein & Marusik, 2012.
Females of R. alpina sp. n., R. gracilis sp. n., R. guangxi, R. rugosa sp. n., R. shangrila, R. songi, R. spirula sp. n. and R. yunnanensis are unknown.
1 | Males | 2 |
– | Females | 15 |
2 | PMS present | 3 |
– | PMS absent | 9 |
3 | Carapace length > 10 mm. Embolus with distinct subapical keel | xizangensis |
– | Carapace length 3.0–7.3 mm. Embolic keel absent or vestigial | 4 |
4 | Palpal tibia relatively short, with a length/width ratio of 3.0–3.2 (Fig. |
montana |
– | Palpal tibia and embolus relatively long, with a length/width ratio of 3.6–4.5 (Figs |
5 |
5 | Embolus with distally hooked tip (Figs |
gracilis sp. n. |
– | Embolic tip not hooked | 6 |
6 | Embolus more or less twisted | 7 |
– | Distal part of embolus curved gradually ( |
hebeinica |
7 | Tibia I equal in length to or shorter than metatarsus (as in Fig. |
8 |
– | Tibia I considerably longer than metatarsus (see |
yunnanensis |
8 | Embolus only slightly twisted (Op. cit.: fig. 42) | songi |
– | Embolus distinctly twisted (Figs |
yajiangensis sp. n. |
9 | Embolus with hooked tip (see |
guangxi |
– | Embolic tip not hooked | 10 |
10 | Embolus strongly spiralled as in Figs |
spirula sp. n. |
– | Embolus more or less curved, slightly spiralled or bent as in Figs |
11 |
11 | Embolus slightly spiralled as in Figs |
alpina sp. n. |
– | Embolus curved or gradually twisted | 12 |
12 | Entire embolus arched as in Figs |
rugosa sp. n. |
– | Embolus more or less distinctly twisted as in Figs |
13 |
13 | Abdomen with dorsal and ventral spotted pattern (Figs |
14 |
– | Abdomen uniformly dark brown. Embolus noticeably twisted (see |
shangrila |
14 | Dorsal abdominal pattern consists of numerous darker spots on a lighter background (Fig. |
bellula sp. n. |
– | Dorsal abdominal pattern consists of numerous lighter spots on a darker background (Fig. |
chayi sp. n. |
15 | PMS present | 16 |
– | PMS absent | 19 |
16 | Carapace length > 10 mm. Median (ental) branch of receptacle bifurcate (Op. cit., fig. 50) | xizangensis |
– | Carapace length < 8 mm. Median (ental) branch of receptacle entire | 17 |
17 | Median (ental) branch of receptacle curved inward as shown in Figs |
yajiangensis sp. n. |
– | Shape of receptacles different | 18 |
18 |
PLS: apical segment triangular. Receptacular bases narrow ( |
hebeinica |
– | PLS: apical segment digitiform. Receptacular bases widened (Op. cit., fig. 48) | montana |
19 | Receptacles as shown in Figs |
bellula sp. n. |
– | Receptacles as shown in Figs |
chayi sp. n. |
Holotype ♂ – CHINA, Yunnan Province, Zhongdian County, northern Zhongdian [27°50.119'N, 99°42.426'E, elevation 3285 m], July 23–30, 2000, X. Yu & H. Zhou (IZCAS). Paratypes: same data but Xiaoxueshanyakou [27°49.119'N, 99°41.426'E, elevation 3265 m] – 1♂ (IZCAS); same data but Bitahaixi [27°48.105'N, 99°40.429'E, elevation 3285 m] – 2♂ (IZCAS).
The specific name is taken from the Latin adjective “alpinus”, which means “alpine” and refers to the high altitude of the type locality.
The new species is similar to R. chayi sp. n., R. shangrila and R. songi, all also occurring in Yunnan, but differs by the slightly twisted and bent distal portion of the embolus (Figs
Male (holotype): TL 14.50, CL 5.75, CW 5.25, AL 6.55, AW 4.00. Eye diameters and interdistances: AME 0.17, ALE 0.31, PME 0.13, PLE 0.15, AME–AME 0.19, AME–ALE 0.10, PME–PME 0.55, PME–PLE 0.08. Leg lengths: I: 17.05 (4.90+2.10+4.40+3.50+2.15), II: 16.15 (4.85+1.50+4.30+3.40+2.10), III: 14.50 (4.25+1.70+3.10+3.25+2.20), IV: 18.45 (5.10+2.15+4.55+4.10+2.55). Labium, sternum and maxillae as shown in Fig.
Female. Unknown.
China: northern Yunnan.
Holotype ♂ – CHINA, Yunnan Province, Mengla County, Xishuangbanna, Menglun Town, primary tropical seasonal rainforest in Menglun Nature Reserve [21°57.445'N, 101°12.997'E, 744 m], January 16–31, 2007, G. Zheng (IZCAS). Paratypes: 26♂, 2♀ (IZCAS), same data as holotype.
The specific name is taken from the Latin adjective “bellulus” (the diminutive form of “bellus”), which means “beautiful” and refers to the perfect shape of the palpal bulb.
This new species is similar to R. chayi sp. n. and R. yunnanensis but can be distinguished by having a considerably shorter embolus than that in R. chayi sp. n. (Figs
Male (holotype): TL 7.90, CL 3.50, CW 2.55, AL 3.60, AW 2.25. Eye diameters and interdistances: AME 0.20, ALE 0.24, PLE 0.16, PME 0.12, AME–AME 0.09, AME–ALE 0.04, PME–PME 0.35, PME–PLE 0.03. Leg lengths: I: 10.04 (2.75+1.65+2.65+1.80+1.55), II: 9.70 (2.75+1.30+2.30+1.85+1.50), III: 9.05 (2.50+1.15+1.65+2.25+1.50), IV: 12.40 (3.25+1.25+3.10+3.15+1.65). Labium, sternum and maxillae as shown in Fig.
Female. TL 6.75, CL 3.25, CW 2.50, AL 2.75, AW 2.40; body and legs colouration as in male. Eye diameters and interdistances: AME 0.22, ALE 0.24, PME 0.10, PLE 0.15, AME–AME 0.08, AME–ALE 0.05, PME–PME 0.36, PME–PLE 0.04, Leg lengths: I: 10.15 (2.65+1.60+2.55+1.90+1.45), II: 9.55 (2.65+1.45+2.25+1.75+1.45), III: 8.75 (2.45+1.10+1.50+2.25+1.45), IV: 12.00 (3.20+1.15+3.00+3.10+1.55). Genital area as in Fig.
Total length of males and females: 6.75–7.93 (n=8).
The species is known only from the type locality.
Holotype ♂ – CHINA: Yunnan Province, Lijiang County, Mt. Yulongxueshan, Maoniuping [27°05.503'N, 100°15.403'E, elevation 3061 m], August 1–3, 2000, X. Yu (IZCAS). Paratypes: 16♂ (IZCAS), same data as holotype; 1♀ – Sichuan Province, Yanyuan County, around Lugu Lake [27°48'N, 100°49'E, elevation 3300 m], May 28, 2011, J. Martens (SMF).
The specific name is from the Chinese word for difference (chā yì), in reference to the difference between the new species with R. songi and R. yunnanensis; noun.
The new species is similar to R. alpina sp. n., R. songi and R. yunnanensis but can be distinguished by the smooth distal portion of the embolus (Figs
Male (holotype): TL 10.30, CL 4.30, CW 3.65, AL 4.90, AW 3.10. Eye diameters and interdistances: AME 0.17, ALE 0.23, PME 0.13, PLE 0.21; AME–AME 0.11, AME–ALE 0.06, PME–PME 0.31, PME–PLE 0.05. Leg lengths: I: 12.50 (3.60+1.25+2.95+2.45+2.25), II: 11.65 (3.55+1.25+2.85+2.50+1.50), III: 10.55 (2.55+1.10+2.50+2.65+1.75), IV: 14.40 (4.00+1.30+3.10+4.05+1.95). Maxillae, labium and sternum as shown in Fig.
Female (paratype): TL 14.75, CL 5.75, CW 4.90, AL 9.00, AW 5.25. Eye diameters and interdistances: AME 0.17, ALE 0.30, PME 0.17, PLE 0.23, AME–AME 0.21, AME–ALE 0.12, PME–PME 0.47, PME–PLE 0.04. Leg lengths: I: 13.15 (4.20+2.25+3.00+2.30+1.40), II: 11.95 (3.70+2.20+2.35+2.30+1.40), III: 11.90 (3.15+2.00+2.05+3.05+1.65), IV: 15.90 (4.25+2.30+3.10+4.30+1.95). Most characters, including the colouration peculiarities (Fig.
Total length of males: 9.49–11.10 (n=8).
China: north-western Yunnan, south-western Sichuan.
Holotype ♂ – CHINA: Zhejiang Province, Hangzhou County, Hangzhou [30°16.276'N, 120°09.178'E, 260 m], July 1980, Z. Chen (IZCAS).
The specific name is taken from the Latin adjective “gracilis”, which means “slender” and refers to the shape of embolus.
This new species can be easily distinguished from all known congeners by its slender and subapically curved embolus (Figs
Male (holotype): TL 8.30, CL 4.05, CW 3.25, AL 3.90, AW 2.40. Eye sizes and interdistances: AME 0.15, ALE 0.22, PME 0.11, PLE 0.15, AME–AME 0.11, AME–ALE 0.07, PME–PME 0.34, PME–PLE 0.04. Leg lengths: I: 11.90 (3.50+1.55+2.95+2.15+1.75), II: 11.00 (3.05+1.60+2.70+1.80+1.85), III: 10.75 (2.75+1.60+2.25+2.25+1.60), IV: 12.85 (3.25+1.60+3.15+3.10+1.75). Carapace yellowish brown dorsally, with a few brownish setae. Eye tubercle blackish brown. Chelicerae reddish dark brown. Sternum, labium, maxillae and legs light brown ventrally. Abdomen dorsally light brown, with blackish cloudy maculae and brownish setae. Ventral surface of abdomen and spinnerets yellowish brown, with dense brownish setae (Fig.
Female. Unknown.
The species is known only from the type locality.
CHINA: Sichuan Province, Baoxing County, Baoxing [30°22.052'N, 102°48.534'E, elevation 1115 m], June 2001, X. Yu & H. Zhou – 3♂ (IZCAS); same county, Qiaoqi Town [30°41.129'N, 102°42.370'E, 2447 m], June 6–7, 1997, leg. Z. Zhang.
This species can be easily distinguished from all known East Asian congeners by its short and stout palpal tibia and by a short and flattened embolus (Fig.
See
Total length: 10.46–11.56 in males from Sichuan (n=8) vs. 15.50 in the male holotype from Yunnan.
China: northern Yunnan and southern Sichuan.
Holotype ♂ – CHINA: Yunnan Province, Lijiang County, Shigu Town, Shigu east [26°52.014'N, 100°13.588'E, elevation 2393 m], July 31 to August 4, 2007, X. Yu (IZCAS).
The specific name is taken from the Latin adjective “rugosus”, meaning “wrinkled” and refers to the wrinkled transition between embolus and bulb.
The new species is similar to R. chayi sp. n. but can be distinguished by its embolus gradually curved to the tip (twisted in the latter species) and by the considerably better developed embolic ridges (Figs
Male (holotype): TL 14.50, CL 6.15, CW 4.65, AL 6.25, AW 4.00. Eye diameters and interdistances: AME 0.20, ALE 0.24, PME 0.17, PLE 0.19, AME–AME 0.09, AME–ALE 0.04, PME–PME 0.36, PME–PLE 0.06. Leg lengths: I: 16.50 (4.60+2.25+4.25+3.10+2.30), II: 13.95 (4.10+1.60+3.55+2.55+2.15), III: 12.70 (3.75+1.25+2.75+2.70+2.25), IV: 16.95 (4.55+1.60+4.25+4.05+2.50). Venter as shown in Fig.
Female. Unknown.
The species is known only from the type locality.
Holotype ♂ – CHINA: Hubei Province, Shennongjia Forest Region, Mt. Guanmenshan [31°25.483'N, 110°21.565'E, elevation 1601 m], July 23–30, 1998, H. Zhou (IZCAS). Paratypes: 22♂ (IZCAS), same data as holotype.
The specific name is taken from the Latin noun “spirula” (the diminutive form of “spira = spiral”), which means “small spiral” and refers to the spiral embolus.
The new species is similar to R. yunnanensis but can be distinguished by a noticeably longer and less spinose cymbium, by its more twisted, corkscrew-shaped distal portion of the embolus (see Fig.
Male (holotype): TL 11.25, CL 3.95, CW 3.50, AL 5.65, AW 2.25. Eye diameters and interdistances: AME 0.11, ALE 0.26, PME 0.09, PLE 0.10, AME–AME 0.08, AME–ALE 0.05, PME–PME 0.28, PME–PLE 0.04. Leg lengths: I: 11.65 (3.25+1.45+3.10+2.30+1.55), II: 10.55 (3.10+1.25+2.55+2.15+1.50), III: 9.65 (2.55+1.05+2.05+2.55+1.75), IV: 13.55 (3.35+1.30+2.75+4.10+2.05). Venter as shown in Fig.
Female. Unknown.
Total length: 10.46–11.56 (n=8).
Known only from the type locality.
Holotype ♂ – CHINA: Sichuan Province, Yajiang County, Yajiang [27°50.119'N, 99°42.426'E, elevation 3285 m], 7 June 2001, X. Yu & H. Zhang (IZCAS). Paratypes: same area but Longjiangbian [27°49.119'N, 99°41.426'E, elevation 3265 m], 27 May 2009, X. Yu & H. Zhang – 1♀ (IZCAS).
The specific epithet, a Latinised adjective, refers to the type locality.
Judging from the shape of the bulb and the distal portion of the embolus, this new species is similar to R. shangrila (
Male (holotype): TL 14.10, CL 5.60, CW 5.45, AL 6.20, AW 3.55. Eye diameters and interdistances: AME 0.19, ALE 0.25, PME 0.16, PLE 0.14, AME–AME 0.15, AME–ALE 0.07, PME–PME 0.45, PME–PLE 0.07. Leg lengths: I: 20.96 (6.09+2.24+5.45+4.49+2.69), II: 24.23 (6.47+3.27+5.83+5.45+3.21), III: 17.76 (4.49+2.18+4.10+4.55+2.44), IV: 22.56 (5.83+2.56+4.81+6.67+2.69). Venter as shown in Fig.
Female (paratype): TL 16.50, CL 6.75, CW 5.80, AL 7.25, AW 5.90. Colouration and most somatic characters as in the male. Eye diameters and interdistances: AME 0.21, ALE 0.35, PME 0.17, PLE 0.23, AME–AME 0.18, AME–ALE 0.12, PME–PME 0.53, PME–PLE 0.07. Leg lengths: I: 16.05 (5.05+2.30+3.90+3.05+1.75), II: 15.15 (4.25+2.75+3.30+2.80+2.05), III: 14.45 (4.30+1.95+2.85+3.25+2.10), IV: 19.90 (5.25+2.25+4.40+5.25+2.75). Receptacles divided into a long digitiform inner branch that is bent inward and a short club-shaped outer lobe that is bent anteriad (Figs
China: southern Sichuan.
Sinopesa maculata Raven & Schwendinger, 1995, by the original designation.
Sinopesa, like Raveniola, differs from Hermacha and Entypesa by lacking serrula and metatarsal preening combs and by possessing two enlarged retroventral spines in males and divided receptacles in females. As in members of Raveniola, the PMS in Sinopesa are reduced in size and even lost in some species – a condition which has never been observed in Hermacha and Entypesa. Sinopesa differs from its close relative Raveniola by a thin and less developed scopula and by the presence of a well-developed male intercheliceral tumescence (which is less developed in Raveniola and completely lost in all Chinese members of this genus).
Six species – S. chengbuensis (Xu & Yin, 2002) (China), S. chinensis (Kulczyński, 1901) (China), S. ninhbinhensis sp. n. (Vietnam), S. kumensis Shimojana & Haupt, 2000 (Ryukyu Isles), S. maculata Raven & Schwendinger, 1995 (Thailand) and S. sinensis (Xu & Yin, 2002) (China). The new species is described below.
Males of S. chengbuensis and females of S. ninhbinhensis sp. n. are unknown
1 | Males | 2 |
– | Females | 6 |
2 | PMS present | 3 |
– | PMS absent | 5 |
3 | Dorsal abdominal pattern present. Palpal tibia cylindrical, embolus hooked | 4 |
– | Abdomen uniformly coloured. Palpal tibia arched, embolus corkscrew-shaped (see |
kumensis |
4 | Large species: TL 17 mm. Embolus long: approximately half as long as palpal tibia (see |
sinensis |
– | Small species: TL 10–12 mm. Embolus short: approximately 0.3 times as long as palpal tibia (see |
chinensis |
5 | Abdomen spotted; embolus corkscrew-shaped ( |
maculata |
– | Abdomen uniformly pale; embolus with hooked tip (Figs |
ninhbinhensis sp. n. |
6 | PMS present; abdomen uniformly coloured; receptacles U- or Y-shaped | 7 |
– |
PMS absent; abdomen spotted; receptacles Y-shaped, with short inner and longer outer branch ( |
maculata |
7 | Receptacles U-shaped, with inner and outer branches equal in length ( |
chengbuensis |
– | Receptacles Y-shaped, with outer branch twisted and much longer than the very short inner branch ( |
kumensis |
Holotype ♂ – VIETNAM: Ninh Binh Province, disturbed forest of Cuc Phuong National Park [20°17.066'N, 105°40.253'E, elevation 273 m], pitfall traps, March 1–30, 2008. Paratypes: 26♂ (IZCAS), same data as holotype.
The specific epithet, a Latinised adjective, refers to the type locality.
The new species shares with S. kumensis the complete absence of an abdominal pattern and the presence of a short male palpal tibia, but it can be distinguished from the latter species by the complete absence of PMS and by the presence of a globular bulb and of a gradually tapering and apically hooked embolus (oval and corkscrew-shaped, respectively) in S. kumensis (see Figs
Male (holotype): TL 7.55, CL 2.95, CW 2.45, AL 3.20, AW 1.85. Eye diameters and interdistances: AME 0.11, ALE 0.14, PME 0.12, PLE 0.10, AME–AME 0.07, AME–ALE 0.04, PME–PME 0.26, PME–PLE 0.03. Leg lengths: I: 9.90 (2.75+1.50+2.55+2.15+1.25), II: 8.10 (2.25+1.15+2.10+1.50+1.10), III: 8.15 (1.75+0.95+2.00+2.15+1.30), IV: 10.60 (2.80+1.15+2.40+2.80+1.45). Venter as shown in Fig.
Female. Unknown.
Total length: 6.95–7.70 (n=10).
Vietnam: Ninh Binh Province.
The authors are grateful to Mr Zuwei Zha for preparing photos of the paper. The manuscript benefitted greatly from comments by the subject editor and the referees Peter Schwendinger and Mykola M. Kovblyuk. Peter Jäger and Julia Altmann kindly helped us to study part of the spider collection of the Senckenberg Museum where one of the paratypes was found. Yuri Marusik helped to prepare a photo of the receptacles of this specimen. English was kindly checked by Sarah Crews. This study was supported by the National Natural Sciences Foundation of China (NSFC-31272280, 31471960) for Shuqiang Li and by the Ministry of Absorption, Israel, for Sergei Zonstein.