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Taxonomic studies on the sac spider genus Clubiona (Araneae, Clubionidae) from Xishuangbanna Rainforest, China
expand article infoJianshuang Zhang, Hao Yu§, Shuqiang Li|
‡ Guizhou Normal University, Guiyang, China
§ Guizhou Education University, Guiyang, China
| Institute of Zoology, Chinese Academy of sciences, Beijing, China
Open Access

Abstract

Spiders of the genus Clubiona Latreille, 1804 from Xishuangbanna, Yunnan Province, China are studied. A total of 47 species is reported and illustrated, including 14 new species and two new synonyms. Twelve of the new species belong to four species groups: C. dengpao Yu & Li, sp. nov., C. subdidentata Yu & Li, sp. nov., C. tixing Yu & Li, sp. nov., C. xiaoci Yu & Li, sp. nov., C. xiaokong Yu & Li, sp. nov., C. yejiei Yu & Li, sp. nov., C. zhaoi Yu & Li, sp. nov. and C. zhigangi Yu & Li, sp. nov. from the C. corticalis group; C. mii Yu & Li, sp. nov. and C. subtongi Yu & Li, sp. nov. from the C. ternatensis group; C. banna Yu & Li, sp. nov. from the C. filicata group; and C. menglun Yu & Li, sp. nov. from the C. trivialis group. The remaining two new species, C. shuangsi Yu & Li, sp. nov. and C. wangchengi Yu & Li, sp. nov., are not readily assignable to any of the existing species groups. The female of C. cochlearis Yu & Li, 2019, the female of C. tiane Yu & Li, 2019, the female of C. bicornis Yu & Li, 2019, the male of C. lala Jäger & Dankittipakul, 2010 and the true female of C. suthepica Dankittipakul, 2008 are described for the first time. Two new synonyms are: C. vukomi Jäger & Dankittipakul, 2010 syn. nov. = C. circulata Zhang & Yin, 1998; C. melanothele Thorell, 1895 syn. nov. = Clubiona melanosticta Thorell, 1890. A checklist of Clubiona species from Xishuangbanna is provided. The DNA barcodes of almost all of the species were obtained for species delimitation, matching of sexes and future use.

Keywords

Checklist, DNA barcoding, new species, new synonymy, taxonomy, tropical rainforest

Introduction

Clubiona Latreille, 1804 is the type genus of the Clubionidae Wagner, 1887 and currently includes 506 extant species that are found worldwide except for the Polar Regions and South America (WSC 2021). This genus comprises 61% of the total number of species of the family (Marusik and Omelko 2018; Zhang and Yu 2020; WSC 2021). Despite its high species diversity, the genus Clubiona remains inadequately studied: almost half of the species are known from a single sex or juveniles (82 from males only, 133 from females only, two from juveniles only), and in some cases, the adults are apparently mismatched, or conspecific males and females have been described as separate species (Deeleman-Reinhold 2001; Jäger and Dankittipakul 2010; WSC 2021); the descriptions from early studies are rather brief, many species are not illustrated, or illustrations are inadequate; types of some species do not exist or are difficult to locate or access.

Clubiona are common spiders in China, with 152 species, of which 106 are known from both sexes (WSC 2021). Except for very few species, almost all of Chinese Clubiona were described or redescribed in the past 30 years (WSC 2021). Clubiona from Thailand and Laos have been well studied by Deeleman-Reinhold (2001), and Dankittipakul and co-authors (Dankittipakul and Singtripop 2008a, b; Dankittipakul et al. 2012; Jäger and Dankittipakul 2010). Clubiona from Myanmar are relatively poorly studied; half of the described species have not been illustrated or descriptions are accompanied by inadequate illustrations (WSC 2021).

Xishuangbanna is a key biogeographic area and a biodiversity hotspot in China (Myers 1988). It shares a border with Myanmar in the southwest and Laos in the southeast and harbours more species diversity than typical tropical rain forests of Southeast Asia (Zhu et al. 2006). Xishuangbanna spiders have received a lot of attention because of an “All Species Inventory” which has been conducted by SL and his team during the last 15 years (Li 2020). Clubiona from the region have been studied by Zhang and Yin (1998), Yin et al. (2012), Wu et al. (2015) and Yu and Li (2019a, b). These studies have described 25 new species and increased the total clubionid species number to 27 in the last 22 years (Yu and Li 2019b). However, based on samples from Xishuangbanna collected during 2007 to 2019, the diversity of the genus Clubiona is underestimated.

In the present paper, a checklist of Xishuangbanna Clubiona spiders is provided based on published literature and new collections. A total of 51 species are recorded, among them, 47 were collected and illustrated, including fourteen new species. The goal of this paper is to provide a detailed description and diagnosis of these new species, to provide the first description of the male or females of 4 known species, to synonymise C. vukomi Jäger & Dankittipakul, 2010 and C. melanothele, Thorell 1895 and to provide comparative illustrations of male palps and female epigynes for all Xishuangbanna Clubiona species.

Materials and methods

Almost all of the species are leaf-dwellers. Most specimens were collected by canopy fogging, while a few were obtained by beating vegetation and pitfall trapping. Specimens were preserved in 75 or 95% ethanol. All type specimens are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing, China (curator Jun Chen).

Specimens were examined using a LEICA M205C and an Olympus SZX7 stereomicroscope. Further details were studied under a CX41 compound microscope. Male and female copulatory organs were examined and illustrated after dissection. Left male palps are illustrated unless otherwise indicated (photos of the right palp were horizontally mirrored in the figures to allow easier comparison with other species). Epigynes were removed and cleared in lactic acid or warm 10% potassium hydroxide (KOH) solution. Some vulvae were imaged after being embedded in Arabic gum. Images were captured with a Canon EOS 70D digital camera 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 taken from the widest distance. The total body length does not include chelicerae or spinnerets. Leg lengths are given as total length (femur, patella + tibia, metatarsus, tarsus). Terminology in the text and figure legends follows Yu and Li (2019a, b), Yu et al. (2017a), Zhang et al. (2018) and Dankittipakul and Singtripop (2008a, b).

A partial fragment (650 bp) of the mitochondrial gene cytochrome c oxidase subunit I (COI) was amplified and sequenced to obtain the genetic distances between morphologically similar species and to confirm identifications and sex pairing accuracy. However, we were unable to obtain good extractions from C. kurosawai Ono, 1986, C. rama Dankittipakul & Singtripop, 2008, C. subyaginumai Yu & Li, 2019, C. tixing sp. nov., C. yejiei sp. nov., and C. zhanggureni Yu & Li, 2019 and the male of C. subquebecana Yu & Li, 2019. Although recorded from Xishuangbanna, the following species were not collected by us and were unavailable for molecular work: C. japonicola Bösenberg & Strand, 1906, C. filoramula Zhang & Yin, 1998, C. heteroducta Zhang & Yin, 1998 and C. zhangyongjingi Li & Blick, 2019.

The primers used were: LCOI1490 (5’-GGTCAACAAATCATAAAGATATTG-3’) and HCOI2198 (5’-TAAACTTCAGGGTGACCAAAAAAT-3’). For additional information on extraction, amplification and sequencing procedures, see Malumbres-Olarte J and Vink (2012). Raw sequences were edited and assembled using BioEdit v.7.2.5 (Hall 1999), and uncorrected pairwise distances between sequences were calculated using MEGA v.10.0 (Tamura et al. 2013). All sequences were analysed using BLAST and are deposited in GenBank. The accession numbers are provided in Table 1.

Table 1.

Checklist of Clubiona species from Xishuangbanna and voucher specimen information.

Species groups Species Sex Figures in the present paper Voucher code GenBank accession number References
milingae group C. yaoi* 62A, B, 72A, B, YHCLU0002 MW731651 Yu and Li 2019a; present paper
80A, 88A, 96A YHCLU0003 MW731650
corticalis group C. cochlearis* 53A, 63A YHCLU0068 MW731626 Yu and Li 2019b; female supplemented in present paper
1, 73A, 81A, 89A YHCLU0079 MW731621
C. dengpao sp. nov.* 2, 73B, 81B, 89B YHCLU0080 MW731620 present paper
C. didentata* 54A, 64A YHCLU0015 MW731648 Zhang and Yin 1998; female supplemented in Yu and Li 2019b; present paper
74E, 82E, 90E YHCLU0016 MW731647
C. kai* 54B, 64B YHCLU0259 MW731584 female supplemented in Yu and Li 2019b; present paper
74C, 82C, 90C YHCLU0052 MW731634
C. kurosawai* 3, 4E, F, 56A, 66A present paper
4A–D, G, H, 76A, 84A, 92A
C. moralis* 5, 6E, F, 55A, 65A, YHCLU0025 MW731643 present paper
6A–D, G, H, 75A, 83A, 91A YHCLU0024 MW731644
C. multidentata* 7, 8E, F, 56B, 66B, YHCLU0076 MW731624 present paper
8A–D, G, H, 75D, 83D, 91D YHCLU0077 MW731623
C. parconcinna* 9, 10E, F, 55C, 65C YHCLU0143 MW731590 present paper
10A–D, G, H, 75C, 83C, 91C YHCLU0260 MW731583
C. pollicaris* 11, 12E, F, 56C, 66C YHCLU0020 MW731646 Wu et al. 2015; present paper
12A–D, G, H, 76B, 84B, 92B YHCLU0021 MW731645
C. rama * 13, 14, 53B, 63B Yu et al. 2017b; present paper
corticalis group C. subdidentata sp. nov.* 15, 74F, 82F, 90F YHCLU0073 MW731625 present paper
C. submoralis* 16, 17E, F, 55B, 65B YHCLU0028 MW731642 Wu et al. 2015; present paper
17A–D, G, H, 75B, 83B, 91B YHCLU0029 MW731641
C. subrama* 53C, 63C YHCLU0083 MW731619 Yu and Li 2019a; present paper
73E, 81E, 89E YHCLU0084 MW731618
C. subyaginumai* 54C, 64C Yu and Li 2019a; present paper
75F, 83F, 91F
C. tixing sp. nov.* 18, 73D, 81D, 89D present paper
C. tiane* 54D, 64D YHCLU0054 MW731632 Yu and Li 2019b; female supplemented in present paper
19, 74D, 82D, 90D YHCLU0053 MW731633
C. xiaoci sp. nov.* 20, 21E, F, 55D, 65D YHCLU0088 MW731614 present paper
21A–D, G, H, 75E, 83E, 91E YHCLU0089 MW731613
C. xiaokong sp. nov.* 22, 74A, 82A, 90A YHCLU0078 MW731622 present paper
C. yejiei sp. nov..* 23, 73C, 81C, 89C present paper
C. zhaoi sp. nov.* 24, 74B, 82B, 90B YHCLU0086 MW731616 present paper
C. zhigangi sp. nov.* 25, 26E, F, 53D, 63D YHCLU0185 MW731586 present paper
26A–D, G, H, 73F, 81F, 89F YHCLU0138 MW731592
ternatensis group C. heteroducta Zhang and Yin 1998
C. mii sp. nov.* 27, 77A, 85A, 93A YHCLU0065 MW731629 present paper
C. subkuu* 57C, 67C YHCLU0039 MW731637 Yu and Li 2019a; present paper
77B, 85B, 93B YHCLU0038 MW731638
C. subtongi sp. nov.* 28, 29, 57D, 67D YHCLU0056 MW731630 present paper
C. theoblicki* 57A, 67A YHCLU0092 MW731612 Yu and Li 2019a; present paper
77C, 85C, 93C YHCLU0093 MW731611
C. tongi* 57B, 67B YHCLU0055 MW731631 Yu and Li 2019b; present paper
77D, 85D, 93D YHCLU0095 MW731610
C. zhengi* 57E, 67E YHCLU0042 MW731636 Yu and Li 2019a; present paper
77E, 85E, 93E YHCLU0043 MW731635
japonicola group C. japonicola ♂♀ Yin et al. 2012
filicata group C. abnormis* 30, 31, 60C, 70C YHCLU0113 MW731597 present paper
C. banan sp. nov.* 32, 33E, F, 58C, 68C YHCLU0104 MW731604 present paper
33A–D, G, H, 78E, 86E, 94E YHCLU0139 MW731591
C. circulata* 34, 35E, F, 59C, 69C YHCLU0108 MW731600 Zhang and Yin 1998; present paper
35A–D, 79A, 87A, 95A YHCLU0156 MW731589
C. reichlini* 36, 37E, F, 58A, 68A YHCLU0263 MW731582 present paper
37A–D, G, H, 79B, 87B, 95B YHCLU0264 MW731581
C. filicata* 38, 39, 58B, 68B YHCLU0107 MW731601 Zhang and Yin 1998; present paper
C. filoramula Zhang and Yin 1998
C. grucollaris* 40, 41E, F, 60A, 70A YHCLU0105 MW731603 present paper
41A–D, G, H, 79C, 87C, 95C YHCLU0106 MW731602
C. lala* 42, 43E, F, 60B, 70B YHCLU0110 MW731599 male supplemented in present paper
43A–D, G, H, 79D, 87D, 95D YHCLU0111 MW731598
C. melanosticta* 44, 45E, F, 59A, 69A YHCLU0011 MW731649 present paper
45A–D, G, H, 78F, 86F, 94F YHCLU0164 MW731588
C. suthepica* 46, 47E, F, 59D, 69D YHCLU0114 MW731596 female supplemented in present paper
47A–D, G, H, 79E, 87E, 95E YHCLU0209 MW731585
C. yueya* 60D, 70D YHCLU0116 MW731595 Yu and Li 2019b; present paper
79F, 87F, 95F YHCLU0117 MW731594
C. zhanggureni* 59B, 69B Yu and Li 2019b; present paper
filicata group C. zhangyongjingi Zhang and Yin 1998; Li and Blick, 2019
trivialis group C. bicornis* 61A, 71A YHCLU0180 MW731587 Yu and Li 2019b; female supplemented in present paper
48, 77F, 85F, 93F YHCLU0099 MW731608
C. cheni* 61B, 71B YHCLU0033 MW731639 Yu and Li 2019a; present paper
78A, 86A, 94A YHCLU0032 MW731640
C. menglun sp. nov.* 49, 78B, 86B, 94B YHCLU0097 MW731609 present paper
C. subasrevida* 61C, 71C YHCLU0100 MW731607 Yu and Li 2019b; present paper
78C, 86C, 94C YHCLU0101 MW731606
C. subquebecana* 61D, 71D Yu and Li 2019a; present paper
78D, 86D, 94D YHCLU0103 MW731605
Species group not assigned C. jiandan* 62C, 72C YHCLU0066 MW731628 Yu and Li 2019b; present paper
80B, 88B, 96B YHCLU0067 MW731627 Yu and Li 2019b; present paper
C. shuangsi sp. nov.* 50, 51E–G, 62D, 72D YHCLU0135 MW731593
51A–D, H, I, 80D, 88D, 96D YHCLU0085 MW731617 present paper
C. wangchengi sp. nov.* 52, 80C, 88C, 96C YHCLU0087 MW731615

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).

Table 2.

List of abbreviations used in the text or figures.

Male palp
conductor
C Note: two types of conductors are considered in the present paper, the first type of conductor is separate from the tegulum, and the second type of conductor is represented by a membranous area fused to the tegulum.
DCA dorsal cymbial apophysis
E embolus
EB embolar base
FA femoral apophysis
PFR prolateral femoral ridge
PPA prolateral patellar apophysis
PTA prolateral tibial apophysis
RPA retrolateral patellar apophysis
RTA retrolateral tibial apophysis
TA tegular apophysis
TH tegular hump
VTA ventral tibial apophysis
Epigyne
A atrium
AAM atrial anterior margin
AM atrial membrane
APM atrial posterior margin
BS bursa
CD copulatory duct
CO copulatory opening
FD fertilisation duct
R epigynal ridge
SB spermathecal base
SH spermathecal head
SP spermatheca
SS spermathecal stalk
Ocular area
AER anterior eye row
ALE anterior lateral eyes
AME anterior median eyes
MOQ median ocular quadrangle
MOQA MOQ anterior width
MOQL length of MOQ
MOQP MOQ posterior width
PER posterior eye row
PLE posterior lateral eyes
PME posterior median eyes
AMEAME distance between AMEs
AMEALE distance between AME and ALE
PMEPME distance between PMEs
PMEPLE distance between PME and PLE
Institutions
IZCAS Institute of Zoology, Chinese Academy of Sciences, Beijing, China
XTBG Xishuangbanna Tropical Botanic Garden, Yunnan, China

Taxonomy

Family Clubionidae Wagner, 1887

Clubiona Latreille, 1804

Clubiona Latreille, 1804: 134 (type species Araneus pallidulus Clerck, 1757).

Hirtia Thorell, 1881: 222 (type species H. ternatensis Thorell, 1891).

Atalia Thorell, 1887: 54 (type species A. concinna Thorell, 1887).

Tolophus Thorell, 1891: 26 (type species T. submaculatus Thorell, 1891).

Paraclubiona Lohmander, 1944: 19 (type species Aranea corticalis Walckenaer, 1802).

Microclubiona Lohmander, 1944: 20 (type species C. trivialis C.L. Koch, 1834).

Hyloclubiona Lohmander, 1944: 20 (subgenus of Microclubiona, type species C. comta C.L. Koch, 1839).

Heteroclubiona Lohmander, 1944: 20 (subgenus of Clubiona, type species C. terrestris Westring, 1851).

Epiclubiona Lohmander, 1944: 20 (subgenus of Clubiona, type species C. neglecta O. Pickard-Cambridge, 1862, not C. similis L. Koch, 1866 as indicated by Wunderlich 2011).

Euryclubiona Lohmander, 1944: 21 (subgenus of Clubiona, type species C. subsultans Thorell, 1875).

Gauroclubiona Lohmander, 1944: 21 (subgenus of Clubiona, type species C. coerulescens L. Koch, 1867).

Bucliona Benoit, 1977: 68 (type species Clubiona dubia O. Pickard-Cambridge, 1869).

Japoniona Mikhailov, 1990: 143 (subgenus of Clubiona, type species C. japonica L. Koch, 1878).

Bicluona Mikhailov, 1994: 52 (subgenus of Clubiona, type species Liocranum jucundum Karsch, 1879).

Marmorclubiona Wunderlich, 2011: 136 (type species C. marmorata L. Koch, 1866).

Breviclubiona Wunderlich, 2011: 139 (type species C. brevipes Blackwall, 1841).

Anaclubiona Ono, 2010: 4 (type species C. zilla Dönitz & Strand, 1906).

Comments

Clubiona sensu lato currently contains more than 500 nominal species and is one of the largest genera of Araneae (Marusik and Omelko 2018; WSC 2021). Several major taxonomic studies on a regional scale have been conducted, e.g., Simon (1932) for the French species, Lohmander (1944) for the Swedish species, Wiehle (1965) for the German species, Wunderlich (2011) for the European species, Edwards (1958) for the North American species, Dondale and Redner (1982) for the Canadian and Alaskan species, Mikhailov (1990, 1991, 1995, 2002, 2012) for the Palaearctic species, and Deeleman-Reinhold (2001) for the Southeast Asian species.

There are 14 generic names that are currently considered junior synonyms of Clubiona (see above list). Beyond that, at least ten subgeneric and 20 species group names have been recognised for subdivisions of the genus (Lohmander 1944; Dondale and Redner 1982; Mikhailov 1990, 1991, 1995, 2002; Deeleman-Reinhold 2001). However, almost all generic and subgeneric statuses were suppressed by Mikhailov (1990, 1991, 1995, 2002, 2012) and Deeleman-Reinhold (2001). At present, only a dozen species group names are used for the taxonomy of the genus. Although there is no agreement on the limits of most species groups, some groups present a distinct set of characters with relatively stable species composition and are currently accepted by most taxonomists. There are at least 16 species groups discussed or frequently used in recent publications: C. apiculata group, C. corticalis group (corresponds to Atalia and Paraclubiona), C. hystrix group (corresponds to Hirtia), C. japonica group (corresponds to Tolophus and Japoniona), C. trivialis group (corresponds to Microclubiona), C. pallidula group and C. obesa group (belongs to Clubiona s. str.), C. abboi group (mainly distributed in North America, may be elevated to genus level in the future), C. similis group (corresponds to Epiclubiona), C. lutescens group (corresponds to Heteroclubiona), C. reclusa group (corresponds to Euryclubiona), C. caerulescens group (corresponds to Gauroclubiona), C. marmorata group (corresponds to Marmorclubiona), C. brevipess group (corresponds to Breviclubiona), C. zilla group (corresponds to Anaclubiona), C. genevensis group.

According to the quite diverse copulatory structures of both sexes, Clubiona sensu lato has been widely regarded as paraphyletic and will likely be split in the future (Wunderlich 2011; Marusik and Omelko 2018; Zhang and Yu 2020). However, we agree with Mikhailov (2012) regarding the need of an extensive, large-scale review of the genus. Consequently, the present study follows the WSC (2021) and Mikhailov (2012), and temporarily places all species in Clubiona sensu lato.

Key to species groups occurring in Xishuangbanna (males)

1 Dorsum of abdomen/carapace/or legs with pattern (Figs 31A, C, 33E, F, 35E, F, 37E, F, 39A, C, 41E, F, 43E, F, 45E, F, 47E, F) C. filicata group
Legs and body dorsally without distinct pattern 2
2 Bulb enlarged or inflated, and protruded or prolapsed, with indistinct sperm duct (Figs 53A–D, 54A–D, 55A–D, 56A–C, 63A–D, 64A–D, 65A–D, 66A–C) C. corticalis group
Tegulum relatively flattened, sperm duct distinct 3
3 Sperm duct simple, U-shaped, or V-shaped in ventral view (Figs 57A–E, 62A, C) 4
Sperm duct sinuous (Figs 61A–D, 62D) 6
4 Conductor absent; embolus relatively long, oriented clockwise along the margin of the tegulum; tegulum distally with a tegular hump (Fig. 57A–E) C. ternatensis group
Conductor present; embolus very short (Figs 62A, C, 72A–C) 5
5 Male palp with a dorsal cymbial apophysis and three tibial apophyses, the retrolateral apophysis well-developed and distally forked (Figs 62B, 72B) C. milingae group (C. yaoi)
Male palp without dorsal cymbial apophysis, tibia only with a simple and not forked retrolateral apophysis (Figs 62C, 72C) C. jiandan
6 Conductor filiform, separated from tegulum (Figs 50C–E, 62D) C. shuangsi sp. nov.
Conductor absent, or depressed and groovelike 7
7 Retrolateral tibial apophysis unbranched (Fig. 71B–D) or bifurcated and processes of different size (Fig. 71A); conductor groovelike, represented by membranous part of tegulum (Figs 61A–D, 71A–D); embolus arched around or angled across the distal end of tegulum, pointed proximally (Figs 61A–D, 71A–D) C. trivialis group
Retrolateral tibial apophysis bifurcated, both processes of similar size; conductor absent; embolus directed retrolaterad, then prolaterodistad C. japonicola group (C. japonicola)

Key to species groups occurring in Xishuangbanna (females)

1 Dorsum of abdomen with pattern (Figs 33G, 35G, 37G, 41G, 43G, 45G, 47G); epigynal atrium broad (almost equal to epigyne width), located at anterior part of epigynal plate (78F, 79A–F, 86F, 87A–F) (atria is relatively small in C. banna sp. nov., Figs 78E, 86E) C. filicata group
Abdomen dorsally without distinct pattern; epigynal atrium absent, or located posteriorly, or located anteriorly but small 2
2 Copulatory openings located at anterior part of epigyne (Figs 81A–F, 82A–F, 83A–F, 84A, 88D) (located in the middle in C. pollicaris, Fig. 84B), well separated from epigastric furrow 3
Copulatory openings located posteriorly, close to epigastric furrow 4
3 Epigynal atrium shaped like an inverted triangle (Figs 80D, 88D) C. shaungsi sp. nov.
Epigynal atrium of variable shapes, but not inverted triangular C. corticalis group
4 Copulatory openings fused, or closely spaced, or separated by no more than one diameter (Figs 77A–F, 78A–D, 85A–F, 86A–D) 5
Copulatory openings separated by more than 1.5 × diameters (Figs 80A–C, 88A–C) 7
5 Copulatory openings hidden in ridges, folds, or under hood (Figs 77A–E, 85A–E) C. ternatensis group
Epigynal ridge or fold absent 6
6 Copulatory openings large, elongate, length more than 1/3 of epigyne length; copulatory ducts as broad as spermathecae C. japonicola group (C. japonicola)
Copulatory openings small, usually circular, diameter no more than 1/5 of epigyne length (Figs 77F, 78A–D, 85F, 86A–D); copulatory ducts slenderer than spermathecae (Figs 93F, 94A–D) C. trivialis group
7 Tibia I with three pairs of ventral spines; bursae oblong (Fig. 96A) C. milingae group (C. yaoi)
Tibia I with two pairs of ventral spines; bursae globular 8
8 Epigynal plate distinctly longer than wide, copulatory openings circular (Figs 80B, 88B), spermathecae oval (Fig. 96B) C. jiandan
Epigynal plate distinctly wider than long, copulatory openings pocket-like (Figs 80C, 88C), spermathecae globular (Fig. 96C) C. wangchengi sp. nov.

Clubiona milingae group

Clubiona apiculata group: Dankittipakul and Singtripop 2014: 1924.

Diagnosis

See Dankittipakul and Singtripop (2014) and Yu and Li (2019a).

Description

See Dankittipakul and Singtripop (2014).

Composition and distribution

Clubiona apiculata Dankittipakul & Singtripop, 2014 (♂♀), C. conica Dankittipakul & Singtripop, 2014 (♂♀), C. cylindriformis Dankittipakul & Singtripop, 2014 (♂) and C. cultrata Dankittipakul & Singtripop, 2014 (♂) endemic to Borneo, C. yaoi Yu & Li, 2019 (♂♀) and C. milingae Barrion-Dupo, Barrion & Heong, 2013 (♂♀) from China.

Comments

The Clubiona milingae group was established by Dankittipakul and Singtripop (2014) for four Borneo species. Yu and co-authors assigned two Chinese species to the species group (Yu and Li 2019; Zhang et al. 2020). The group presents a distinct set of characters, can be considered monophyletic, and may deserve the status of a separate genus in the future.

Clubiona yaoi Yu & Li, 2019

Figs 62A, B, 72A, B, 80A, 88A, 96A

Clubiona yaoi Yu & Li, 2019a: 152, figs 1A–E, 2A–H (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34498), 1♀ (paratype, IZCAS Ar 34499), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary forest, 21°54.459'N, 101°16.755'E, ca. 644 m, 20.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♂ (YHCLU0002), XTBG, leprosy village, 21°53.593'N, 101°17.329'E, ca. 559 m, 5.VIII.2018, H. Yu et al. leg.; 1♀ (YHCLU0003), XTBG, teak plantation, 21°54.117'N, 101°16.167'E, ca. 549 m, 8.VIII.2018, H. Yu et al. leg.

Diagnosis and description

See Yu and Li (2019a). Male palp as in Figs 62A, B, 72A, B, epigyne as in Figs 80A, 88A, 96A.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubuiona milingae.

Clubiona corticalis group

Atalia Thorell, 1887: 54 (type species A. concinna Thorell, 1887).

Clubiona: Simon 1897: 76 (synonymised Atalia); Deeleman-Reinhold 2001: 90 (synonymised Paraclubiona).

Clubiona corticalis group: Simon 1932: 905; Mikhailov 1990: 142; Deeleman-Reinhold 2001: 90.

Paraclubiona Lohmander, 1944: 19 (type species Aranea corticalis Walckenaer, 1802).

Diagnosis

See Mikhailov (1995), Deeleman-Reinhold (2001), and Yu and Li (2019a).

Description

See Mikhailov (1995) and Deeleman-Reinhold (2001).

Composition and distribution

Based on previous publications (Mikhailov 1995, 1998; Deeleman-Reinhold 2001; Ono and Hayashi 2009; Huang and Chen 2012; Wu et al. 2015; Yu and Li 2019a, b; Zhang and Yu 2020), at least 67 Clubiona species have been assigned to the corticalis group (from rows 1–67 in Table 3). A few other known species (from rows 68–73 in Table 3) resemble to some species in rows 1–67, but as no one indicated the group placement of these species, they are assigned tentatively to the corticalis group in the present paper for the lack of a better solution.

Comments

At least two generic names are available for the corticalis group, Atalia Thorell, 1887 (type species A. concinna) and Paraclubiona Lohmander, 1944 (type species C. corticalis) (Zhang et al. 2018). The two taxa are currently considered junior synonyms of Clubiona (Wu et al. 2015; WSC 2021). The corticalis group is one of the most speciose clubionid groups and can be further divided into at least four or five subgroups based on morphological characters and molecular data (pers. obs.). We believe that the group deserves the status of a separate genus that can be further divided into several species groups in the future. A review of the genus Clubiona sensu lato and the corticalis group are not within the scope of this work.

Most species of the C. corticalis group are known from both sexes (Table 3); six species are known from males only: C. fanjingshan, C. huiming, and C. subcylindrica from Mt. Fanjing in Guizhou Province (1000 km from Xishuangbanna) and C. lamina, C. aculeata, and C. tengchong from northwest Yunnan (ca. 500 km from Xishuangbanna, Southeast Yunnan). We describe six new species known from females only in the present paper. In consideration of limited distribution ranges in almost all of the corticalis group species (Table 3), Xishuangbanna species are less likely to conspecific to the six species which are known from males only. None of our new species could be matched with C. lamina, C. aculeata, and C. tengchong due to their different habitus: Xishuangbanna species exhibit typical Southeast Asian corticalis group features, such as a lack of dark markings on the abdomen (Figs 2F, G, 15F, G, 18F, G, 22F, G, 23F, G, 24F, G) (vs. posteriorly with several chevron-shaped patterns dorsally on the abdomen of C. lamina, C. aculeata, and C. tengchong).

Table 3.

Clubiona corticalis group species.

Species name Known sex3 Distribution
1 C. aculeata Zhang, Zhu & Song, 2007 China (Yunnan)
2 C. allotorta Dankittipakul & Singtripop, 2008 ♂♀ Thailand (Chiang Mai)
3 C. alticola Dankittipakul & Singtripop, 2008 ♂♀ Thailand (Chiang Mai)
4 C. altissimoides Liu, Yan, Griswold & Ubick, 2007 ♂♀ China (Yunnan)
5 C. altissimus Hu, 2001 China (Xizang)
6 C. applanata Liu, Yan, Griswold & Ubick, 2007 ♂♀ China (Yunnan)
7 C. bandoi Hayashi, 1995 ♂♀ Japan (Shikoku)
8 C. biforamina Liu, Peng & Yan, 2016 ♂♀ China (Yunnan)
9 C. bifurcata Zhang, Yu & Zhong, 2018 ♂♀ China (Guizhou)
10 C. bomiensis Zhang & Zhu, 2009 ♂♀ China (Xizang)
11 C. boxaensis Biswas & Biswas ♂♀ India (Jalpaiguri)
12 C. brachyptera Zhu & Chen, 2012 ♂♀ China (Hainan)
13 C. caohai Zhang & Yu, 2020 ♂♀ China (Guizhou)
14 C. chakrabartei Majumder & Tikader, 1991 India (Uttarakhand)
15 C. cirrosa Ono, 1989 ♂♀ Japan (Ryukyu Is.)
16 C. cochlearis Yu & Li, 2019 ♂♀ China (Yunnan)
17 C. cochleata Wang, Wu & Zhang, 2015 ♂♀ China (Yunnan)
18 C. concinna (Thorell, 1887) ♂♀ Myanmar (Tharrawaddy)
19 C. cordata Zhang & Zhu, 2009 ♂♀ China (Sichuan, Xizang)
20 C. corticalis (Walckenaer, 1802) ♂♀ Europe, Turkey, Caucasus
21 C. cylindrata Liu, Yan, Griswold & Ubick, 2007 ♂♀ China (Yunnan)
22 C. dactylina Liu, Peng & Yan, 2016 ♂♀ China (Yunnan)
23 C. dakong Zhang & Yu, 2020 China (Xiang)
24 C. dichotoma Wang, Chen & Z.S. Zhang, 2018 ♂♀ China (Guizhou)
25 C. didentata Zhang & Yin, 1998 ♂♀ China (Yunnan)
26 C. falciforma Liu, Peng & Yan, 2016 ♂♀ China (Yunnan)
27 C. fanjingshan Wang, Chen & Z.S. Zhang, 2018 China (Guizhou)
28 C. femorocalcarata Huang & Chen, 2012 ♂♀ China (Taiwan)
29 C. globosa Wang, Chen & Z. S. Zhang, 2018 ♂♀ China (Guizhou)
30 C. gongshan He, Liu & Zhang, 2016 ♂♀ China (Yunnan)
31 C. huiming Wang, F. Zhang & Z. S. Zhang, 2018 China (Guizhou)
32 C. kai Jäger & Dankittipakul, 2010 ♂♀ Laos (Luang Prabang), China (Yunnan)
33 C. kasanensis Paik, 1990 ♂♀ Korea (Gangwon, Gyeongsangbuk, Jeollabuk), Japan (Kojima)
34 C. kayashimai Ono, 1994 China (Taiwan)
35 C. kuanshanensis Ono, 1994 China (Taiwan)
36 C. kurosawai Ono, 1986 China, Korea, Japan
37 C. lamellaris Zhang, Yu & Zhong, 2018 ♂♀ China (Guizhou)
38 C. lamina Zhang, Zhu & Song, 2007 China (Yunnan)
39 C. lucida He, Liu & Zhang, 2016 ♂♀ China (Hunan)
40 C. lyriformis Song & Zhu, 1991 China (Hubei)
41 C. medog Zhang, Zhu & Song, 2007 China (Xizang)
42 C. mikhailovi Deeleman-Reinhold, 2001 Indonesia (Java)
43 C. moralis Song & Zhu, 1991 ♂♀ China (Yunnan, Hubei, Taiwan)
44 C. multidentata Liu, Peng & Yan, 2016 ♂♀ China (Yunnan)
45 C. parallela Hu & Li, 1987 ♂♀ China (Xizang)
46 C. parconcinna Deeleman-Reinhold, 2001 ♂♀ Thailand (Nakhon Ratchasima), Indonesia (Borneo), China (Yunnan).
47 C. pianmaensis Wang, Wu & Zhang, 2015 ♂♀ China (Yunnan)
48 C. pollicaris Wu, Zheng & Zhang, 2015 ♂♀ China (Yunnan)
49 C. pototanensis Barrion & Litsinger, 1995 Philippines (Panay Is.)
50 C. pyrifera Schenkel, 1936 ♂♀ China (Gansu, Hubei)
51 C. qiyunensis Xu, Yang & Song, 2003 ♂♀ China (Fujian, Anhui)
52 C. rama Dankittipakul & Singtripop, 2008 ♂♀ India (Wes Bengal), Thailand (Phitsanulok), China (Yunnan)
53 C. ryukyuensis Ono, 1989 ♂♀ Japan (Ryukyu Is.)
54 C. stiligera Deeleman-Reinhold, 2001 ♂♀ Indonesia (Sumatra)
55 C. subapplanata Wang, Chen & Z.S. Zhang, 2018 ♂♀ China (Guizhou)
56 C. subcylindrica Wang, Chen & Z.S. Zhang, 2018 China (Guizhou)
57 C. submoralis Wu, Zheng & Zhang, 2015 ♂♀ China (Yunnan)
58 C. subrama Yu & Li, 2019 ♂♀ China (Yunnan)
59 C. subyaginumai Yu & Li, 2019 ♂♀ China (Yunnan)
60 C. taiwanica Ono, 1994 ♂♀ China (Taiwan)
61 C. tangi Liu, Peng & Yan, 2016 ♂♀ China (Yunnan)
62 C. tengchong Zhang, Zhu & Song, 2007 China (Yunnan)
63 C. tiane Yu & Li, 2019 ♂♀ China (Yunnan)
64 C. tortuosa Zhang & Yin, 1998 China (Yunnan)
65 C. violaceovittata Schenkel, 1936 China (Gansu)
66 C. yaginumai Hayashi, 1989 ♂♀ China (Taiwan), Japan (Honshu)
67 C. yanzhii Zhang & Yu, 2020 China (Hunan)
68 C. bucera Yang, Ma & Zhang, 2011 ♂♀ China (Yunnan)
69 C. linzhiensis Hu, 2001 ♂♀ China (Xizang)
70 C. ovalis Zhang, 1991 China (Fujian)
71 C. pseudocordata Dhali, Roy, Saha & Raychaudhuri, 2016 India (West Bengal)
72 C. wolongica Zhu & An, 1999 ♂♀ China (Anhui)
73 C. zhangmuensis Hu & Li, 1987 ♂♀ China (Xizang)
74 C. dengpao Yu & Li, sp. nov. China (Yunnan)
75 C. subdientata Yu & Li, sp. nov. China (Yunnan)
76 C. tixingi Yu & Li, sp. nov. China (Yunnan)
77 C. xiaoci Yu & Li, sp. nov. ♂♀ China (Yunnan)
78 C. xiaokong Yu & Li, sp. nov. China (Yunnan)
79 C. yejiei Yu & Li, sp. nov. China (Yunnan)
80 C. zhaoi Yu & Li, sp. nov. China (Yunnan)
81 C. zhigangi Yu & Li, sp. nov. ♂♀ China (Yunnan)

Key to C. corticalis group species occurring in Xishuangbanna (males)

Males of C. dengpao sp. nov., C. subdidentata sp. nov., C. tixing sp. nov., C. xiaokong sp. nov., C. yejiei sp. nov. and C. zhaoi sp. nov. are excluded due to a lack of specimens.

1 Palp with femoral apophysis (Figs 56C, 66C) C. pollicaris
Palpal femur unmodified 2
2 Palp with patellar apophysis (Figs 65A–D, 66B) 3
Palpal patella unmodified 7
3 Palpal tibia retrolaterally with several short, modified spines near the base (Figs 55D, 65C, D) 4
Palpal tibia without spines 5
4 Conductor absent; retrolateral patellar apophysis with short, modified spines (Figs 55D, 65D) C. xiaoci sp. nov.
Conductor distinct; retrolateral patellar apophysis without short, modified spines (Figs 55C, 65C) C. parconcinna
5 Patellar retrolateral apophysis represented by a small conoid, patella with a row of longitudinally arranged teeth in retrolateral view (Fig. 66B) C. multidentata
Patellar retrolateral apophysis with an indented tip; patella retrolaterally without small tooth 6
6 Embolus wide and triangular (Fig. 55B) C. submoralis
Embolus claw-like and curved (Fig. 55A) C. moralis
7 Palpal tibia with single retrolateral apophysis (Figs 64A–D, 66A) 8
Palpal tibia with 2 retrolateral apophyses (Fig. 63A–D) 12
8 Bulb proximally with an apophysis (Fig. 56A); retrolateral tibial apophysis with thin distally and wide basally (Fig. 66A) C. kurosawai
Bulb proximally without apophysis (Fig. 54A–D); retrolateral tibial apophysis not subdivided (Fig. 64A–D) 9
9 Both conductor and tegular apophysis present (Figs 54A, B, 64A, B) 10
Both conductor and tegular apophysis absent (Figs 54C, D, 64C, D) 11
10 Embolus twisted, distinctly longer than conductor; conductor papilliform with membranous tip; tegular apophysis distinctly smaller than embolus, tooth-shaped, directed prolatero-distally (Figs 54B, 64B) C. kai
Embolus slightly curved, approximately as long as conductor; conductor linguiform and heavily sclerotised; tegular apophysis almost the same size as embolus, triangular, directed distally (Figs 54A, 64A) C. didentata
11 Embolus curved and neck of a swan-shaped (Fig. 54D); retrolateral tibial apophysis shaped like dorsal fin of a fish (Fig. 64D) C. tiane
Embolus straight and needle-shaped (Fig. 54C); retrolateral tibial apophysis thumb-like (Fig. 64C) C. subyaginumai
12 Embolus strong, spoon-shaped, with expanded, torsional tip (Figs 53A, 63A) C. cochlearis
Embolus slender and filiform (Figs 53B–D, 63B–D) 13
13 Embolar apex sinuate; conductor short, ca. 1/5 of tegulum length, with a blunt tip (Figs 53D, 63D) C. zhigangi sp. nov.
Embolar tip not curved; conductor long, not less than 1/3 of tegulum length, with a sharply pointed tip (Figs 53B, C, 63B, C) 14
14 Retrolateral tibial apophysis with sharp apex; ventral tibial apophysis trapezoidal, with blunt tip (Fig. 63C) C. subrama
Retrolateral tibial apophysis apically indented; ventral tibial apophysis subtriangular, with sharp tip (Fig. 63B) C. rama

Key to C. corticalis group species occurring in Xishuangbanna (females)

C. rama is excluded due to lack of specimens.

1 Copulatory openings located in the centre of epigynal plate (Figs 76B, 84B) C. pollicaris
Copulatory openings located at anterior part of epigynal plate 2
2 Spermathecae tubular and sinuous, spermatheca with head (Figs 89A–F, 90A, D, 92A) 3
Spermathecae not as above 11
3 Atrium not less than 1/3 of epigyne width (Figs 73A–D, 81A–D) 4
Atrium reduced, relatively small, less than 1/3 of epigyne width (Figs 73E, F, 74A, D, 76A, 81E, F, 82A, D, 84A) 7
4 Atrium with atrial membrane on anterior margin (Figs 73D, 81D) C. tixing sp. nov.
Atrium without atrial membrane (Figs 73A–C, 81A–C) 5
5 Copulatory openings large, diameter as long as atrium length, situated laterally in atrium (Figs 73A, 81A); copulatory ducts and spermathecae thin and strongly twisted (Fig. 89A) C. cochlearis
Copulatory openings small, diameter ca. 1/3 of atrium length, located posteriorly in atrium (Figs 73B, C, 81B, C); copulatory ducts and spermathecae thick and slightly twisted (Fig. 89B, C) 6
6 Atrium light bulb-shaped (Figs 73B, 81B) C. dengpao sp. nov.
Atrium ellipsoid (Figs 73C, 81C) C. yejiei sp. nov.
7 Atrium reduced (Figs 76A, 84A); bursae globular (Fig. 92A) C. kurosawai
Atrium present (Figs 73E, F, 74A, D, 81E, F, 82A, D); bursae ovoid or reniform 8
8 Atrium narrowed, anteriorly cordiform, posteriorly elongate (Figs 73E, F, 81E, F) 9
Atrium not as above 10
9 Copulatory ducts distinctly long, with a long course forming 2 loops before entering spermathecae (Fig. 89E) C. subrama
Copulatory duct relatively short, directly connected to spermathecae (Fig. 89F) C. zhigangi sp. nov.
10 Copulatory ducts thick, heavily sclerotised (Fig. 90A) C. xiakong sp. nov.
Copulatory duct indistinct, almost invisible in dorsal view (Fig. 90D) C. tiane
11 Atrium with atrial membrane on anterior margin (Figs 74C, E, F, 82 C, E, F) 12
Atrial membrane absent (Figs 74B, 75A–F, 82B, 83A–F) 14
12 Atrial membrane disc-shaped (Figs 74C, 82C); bursae sclerotised (Fig. 90C) C. kai
Atrial membrane subtriangular (Figs 74E, F, 82E, F); bursae membranous (Fig. 90E, F) 13
13 Atrial membrane tongue-shaped (Figs 74E, 82E); copulatory ducts absent (Fig. 90E) C. didentata
Atrial membrane equilateral triangle (Figs 74F, 82F); copulatory ducts present (Fig. 90F) C. subdidentata sp. nov.
14 Copulatory openings widely separated by ca. 2.5–3.0 diameters (Figs 75D, 83D) C. multidentata
Copulatory openings partly fused or close together, separated by not more than one diameter (Figs 74B, 75A–C, E, F, 82B, 83A–C, E, F) 15
15 Spermathecae subtriangular (Figs 90B, 91F) 16
Spermathecae almost spherical (Fig. 91A–C, E) 17
16 Atrial anterior margin M-shaped (Figs 74B, 82B); copulatory ducts indistinct (Fig. 90B) C. zhaoi sp. nov.
Atrial anterior margin transverse (Figs 75F, 83F); copulatory ducts longer than spermathecae diameter (Fig. 91F) C. subyaginumai
17 Epigynal plate anteriorly with a nearly horizontal K-shaped sclerite (Fig. 75A, B) 18
Epigynal plate not as above; atrial anterior margin cambered (Fig. 75C, E) 19
18 Copulatory openings separated by ca. one diameter (Figs 75A, 83A); copulatory ducts descend obliquely (Fig. 91A); bursae 2 × longer than the spermathecae (Fig. 91A) C. moralis
Copulatory openings close together (Figs 75B, 83B); copulatory ducts descend longitudinally (Fig. 91B); spermathecae almost as large as bursae (Fig. 91B) C. submoralis
19 Copulatory openings separated (Figs 75C, 83C); copulatory ducts thin, diameter ca. 1/4 of spermathecae, descend longitudinally (Fig. 91C); bursae surface relatively smooth (Fig. 91C) C. parconcinna
Copulatory openings partly fused (Figs 75E, 83E); copulatory ducts thick, ca. the same diameter as spermathecae, descend obliquely (Fig. 91E); bursae surface wrinkled (Fig. 91E) C. xiaoci sp. nov.

Clubiona cochlearis Yu & Li, 2019

Figs 1, 53A, 63A, 73A, 81A, 89A

Clubiona cochlearis Yu & Li, 2019b: 202, figs 1A–E, 2A–C (♂).

Material examined

Types. Holotype ♂ (IZCAS Ar 34701), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Paramichelia baillonii plantation, 21°54.772'N, 101°16.043'E, ca. 556 m, 18.VII.2007, G. Zheng leg. Other material examined. 1♀, Jinghong City: Pingguan Town, monsoon forest, 22°13.212'N, 100°53.151'E, ca. 832 m, 21.VII.2012, Q.Y. Zhao and C.X. Gao leg; 1♂ (YHCLU0068), Mengyang County: monsoon forest, 21°54.117'N, 101°55.210'E, ca. 856 m, 16.VII.2012, Q.Y. Zhao and C.X. Gao leg; 1♀ (YHCLU0079), Jinghong City: Pingguan Town, monsoon forest, 22°13.668'N, 100°53.351'E, ca. 888 m, 20.VII.2012, Q.Y. Zhao and C.X. Gao leg.

Diagnosis

Females of C. cochlearis are similar to those of C. lyriformis (Yin et al. 2012: 1110, fig. 586a–c). The two species share a similarly large atrium, tubular and sinuous spermathecae, and the copulatory ducts are proximally wide and distally narrow. They differ in the following: (1) atrium is nearly apple-shaped in C. cochlearis (Figs 1A–C, 73A, 81A) (vs. atrium shaped like a violin in C. lyriformis; Yin et al. 2012: fig. 586b); (2) size and location of copulatory openings (copulatory openings are larger and situated laterally in atrium in C. cochlearis vs. relatively small and located at basolateral atrial borders in C. lyriformis) (cf. Figs 1A–C, 73A, 81A and Yin et al. 2012: fig. 586c). Males of C. cochlearis can be easily recognised by the robust, spoon-shaped embolus (Figs 53A, 63A) from all others in the species group.

Description

Male. See Yu and Li (2019b). Male palp as in Figs 53A, 63A.

Female (Fig. 1F, G). Total length 7.38; carapace 3.64 long, 2.61 wide; opisthosoma 3.75 long, 2.53 wide. Carapace brown, darker in cephalic area, without distinct pattern; cephalic region slightly narrowed, cervical groove indistinct; tegument smooth, with short, fine setae. Eyes: AER slightly recurved, PER slightly wider than AER, almost straight in dorsal view. Eye sizes and interdistances: AME 0.16, ALE 0.10, PME 0.13, PLE 0.15, AMEAME 0.21, AMEALE 0.18, PMEPME 0.49, PMEPLE 0.38, MOQL 0.40, MOQA 0.49, MOQP 0.76. Chelicerae robust and brownish red, with four promarginal teeth and two retromarginal denticles. Sternum pale brown, 1.86 long, 1.24 wide. Labium and endites coloured as carapace. Legs light brown, without distinct markings. Leg measurements: I 7.18 (2.20, 2.88, 1.33, 0.77), II 7.66 (12.21, 3.12, 1.47, 0.85), III 7.11 (2.34, 2.53, 1.60, 0.65), IV 9.65 (2.93, 3.32, 2.61, 0.78). Abdomen oval, dorsally grey with dense setae and a lengthwise white heart mark reaching posterior half; with a pair of muscle depressions located at distal part of heart mark; venter uniformly white, without pattern.

Epigyne (Figs 1A–E, 73A, 81A, 89A). Epigynal plate nearly as broad as long, spermathecae and bursae indistinctly visible through integument. Atrium ca. 1/3 of epigyne length and width, more or less apple-shaped, slightly concave anteromedially. Copulatory openings large, located at lateral atrial borders. Copulatory ducts long, proximally thick-walled, extend posteriorly, the latter half slender, ascending obliquely, then connecting with spermathecae at central axis of the vulva. Spermathecae tubular, long and sinuous, strongly convoluted. Fertilisation ducts short and curved, acicular. Bursae situated posteriorly, ovoid, relatively large, close together, ca. 1.5 × longer than wide, surface translucent and smooth.

Distribution

Known only from Xishuangbanna.

Remarks

The female of the species is described for the first time.

Clubiona dengpao Yu & Li, sp. nov.

Figs 2, 73B, 81B, 89B

Holotype

♀ (IZCAS-Ar 34748), China: Yunnan Province: Xishuangbanna: Jinghong City: Menga Town: Wengnan Village: secondary forest, 21°24.265'N, 101°37.296'E, ca. 693 m, 28.VI.2012, Q.Y. Zhao and C.X. Gao leg. Other material examined. 1♀ (YHCLU0080), Mengla County: Nanshahe Village: monsoon forest, 21°36.200'N, 101°34.384'E, ca. 826 m, 14.VII.2012, Q.Y. Zhao and C.X. Gao leg.

Etymology

The specific name is derived from the Chinese pinyin dēng pào, which means ‘lamp bulb’, referring to the atrium which is shaped like a light bulb; noun in apposition.

Diagnosis

This new species is similar to C. cochlearis (Figs 1A–E, 73A, 81A, 89A) and C. yejiei sp. nov. (Figs 23A–E, 73C, 81C, 89C) in having a large atrium and long, tubular and sinuous spermathecae but can be easily distinguished by the light bulb-shaped atrium (Figs 2A–C, 73B, 81B) (vs. apple-shaped in C. cochlearis and ellipsoid in C. yejiei; Figs 1A–C, 73A, 81A, 23A–C, 73C, 81C), and by lacking a fovea (Fig. 2F) (vs. fovea present in C. cochlearis and C. yejiei; Figs 1F, 23F).

Description

Female. Holotype (Fig. 2F, G): Total length 7.86; carapace 3.98 long, 2.51 wide; opisthosoma 3.87 long, 2.36 wide. Carapace greyish white, without pattern, fovea absent; pars cephalica slightly narrowed, cervical groove and radial grooves indistinguishable; tegument smooth, marginally clothed with dense setae. Eyes: AER slightly recurved, PER slightly wider than AER, almost straight in dorsal view. Eye sizes and interdistances: AME 0.11, ALE 0.19, PME 0.18, PLE 0.14, AMEAME 0.18, AMEALE 0.15, PMEPME 0.44, PMEPLE 0.28, MOQL 0.52, MOQA 0.49, MOQP 0.79. Chelicerae coloured as carapace, with four teeth on promargin and three teeth on retromargin. Sternum nearly pure white, 1.99 long, 1.15 wide. Labium and endites coloured as carapace, sparsely covered with setae. Legs coloured as carapace, dorsally covered with black setae, without markings. Leg measurements: I 6.85 (2.01, 2.67, 1.33, 0.85), II 7.21 (2.19, 2.82, 1.45, 0.75), III 6.57 (2.07, 2.22, 1.64, 0.65), IV 8.94 (2.65, 2.79, 2.58, 0.93). Abdomen oval, dorsum light grey, anteriorly and marginally with conspicuous tuft of brown setae, heart mark and muscle depressions indistinct; venter coloured as dorsum, without pattern.

Epigyne (Figs 2A–E, 73B, 81B, 89B). Epigynal plate slightly longer than wide, margin not rebordered; copulatory ducts, spermathecae and bursae indistinctly visible through transparent integument. Atrium large, shaped like a tungsten lamp bulb, ca. 1.2 × longer than wide, ca. 1/3 of epigyne length. Copulatory openings indistinct, close together, located at posterior atrial margin. Copulatory ducts short and thick, heavily sclerotised, descend obliquely, then connect to spermathecae. Spermathecae long and wrinkled, consisting of tubular proximal part and sac-like distal part, with thin fertilisation ducts terminally. Bursae large, ovoid, 1.3 × longer than wide, surface translucent and wrinkled.

Male. Unknown.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona didentata Zhang & Yin, 1998

Figs 54A, 64A, 74E, 82E, 90E

Clubiona didentata Zhang & Yin, 1998: 11, figs 6–8 (♂); Yu and Li 2019b: 207, figs 5A–E, 6A–G (♂♀).

Material examined

1♂ (YHCLU0015), 1♀ (YHCLU0016), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, flower garden, 21°55.915'N, 101°15.099'E, ca. 550 m, 18.VII.2018, X.Q Mi et al. leg; 1♂, XTBG, low evergreen forest, 21°53.794'N, 101°17.152'E, ca. 594 m, 27.XI.2009, G. Tang and Z.Y. Yao leg; 2♀, XTBG, bamboo plantation, 21°53.640'N, 101°16.940'E, ca. 580 m, 3.XII.2009, G. Tang and Z.Y. Yao leg; 1♂3♀, XTBG, rubber-tea plantation, 21°55.239'N, 101°15.854'E, ca. 572 m, 28.VII.2018, Z.G. Chen et al. leg.

Diagnosis and description

See Yu and Li (2019b). Male palp as in Figs 54A, 64A, epigyne as in Figs 74E, 82E, 90E.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona kai.

Clubiona kai Jäger & Dankittipakul, 2010

Figs 54B, 64B, 74C, 82C, 90C

Clubiona kai Jäger & Dankittipakul, 2010: 25, figs 4–12 (♂); Yu and Li 2019b: 207, figs 7A–E, 8A–G (♂♀).

Material examined

1♀ (YHCLU0052), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, 300 acre-feet teak plantation, 21°54.033'N, 101°16.400'E, ca. 554 m, 10.VIII.2018, Z.G. Chen et al. leg; 1♂ (YHCLU0259), XTBG, Anogeissus acuminata plantation, 21°53.992'N, 101°16.948'E, ca. 596 m, 9.V.2019, Z.L. Bai et al. leg; 1♂, XTBG, Lvshilin Forest Park, limestone tropical seasonal rainforest, 21°54.714'N, 101°16.953'E, ca. 660 m, 16.XI.2009, G. Tang and Z.Y. Yao leg; 3♂7♀, XTBG, Lvshilin Forest Park, limestone tropical seasonal rainforest, 21°54.555'N, 101°16.860'E, ca. 610 m, 29.XI.2009, G. Tang and Z.Y. Yao leg.

Diagnosis and description

See Yu and Li (2019b). Male palp as in Figs 54B, 64B, epigyne as in Figs 74C, 82C, 90C.

Distribution

Laos (Luang Prabang), China (Yunnan).

Most similar species

Clubiona didentata.

Clubiona kurosawai Ono, 1986

Figs 3, 4, 56A, 66A, 76A, 84A, 92A

Clubiona kurosawai Ono, 1986: 20, figs 1–8 (♂♀); Mikhailov 1995: 34, fig. 3 (♂); Huang and Chen 2012: 77, figs 22A–F, pl. 6C–D, 7A–B, box 2D (♂♀). For full list of taxonomic references see WSC (2021).

Material examined

1♂, 1♀, China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, rubber plantation, 21°54.350'N, 101°16.461'E, ca. 614 m, 11.VIII.2007, G. Zheng leg.

Diagnosis and description

See Ono (1986). Male palp as in Figs 3, 56A, 66A, epigyne as in Figs 4A–D, 66A, 84A, 92A, habitus as in Fig. 4E–H.

Distribution

Korea, Japan (from Honshu to Southwest Islands), China (Yunnan, Taiwan).

Most similar species

Clubiona bucera.

Clubiona moralis Song & Zhu, 1991

Figs 5, 6, 55A, 65A, 75A, 83A, 91A

Clubiona moralis Song & Zhu, in Song et al. 1991: 70, fig. 5A–D (♂♀); Song and Li 1997: 429, fig. 24A–D (♂♀); Song et al 1999: 426, figs 250M–N, 252Q–R (♂♀); Huang and Chen 2012: 80, fig. 23A–F, pl. 7C (♂♀).

Material examined

1♂ (YHCLU0025), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Paramichelia baillonii plantation, 21°54.183'N, 101°16.967'E, ca. 596 m, 1.VIII.2018, Z.G. Chen leg; 1♀ (YHCLU0024), XTBG, secondary tropical forest, 21°54.833'N, 101°16.781'E, ca. 575 m, 31.VII.2018, Z.G. Chen leg; 4♂6♀, XTBG, Anogeissus acuminata plantation, 21°53.992'N, 101°16.948'E, ca. 596 m, 2.XII.2009, G. Tang and Z.Y. Yao leg.

Diagnosis and description

See Huang and Chen (2012). Male palp as in Figs 5, 55A, 65A, epigyne as in Figs 6A–D, 75A, 83A, 91A, habitus as in Fig. 6E–H.

Distribution

China (Yunnan, Hubei, Taiwan).

Most similar species

Clubiona submoralis.

Clubiona multidentata Liu, Peng & Yan, 2016

Figs 7, 8, 56B, 66B, 75D, 83D, 91D

Clubiona multidentata Liu et al., 2016: 569, figs 37–50 (♂♀).

Material examined

1♂ (YHCLU0076), 1♀ (YHCLU0077), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, 48th km landmark in Menglun Nature Reserve, 21°58.704'N, 101°19.748'E, ca. 1088 m, 11.VIII.2011, G. Zheng et al. leg; 2♂5♀, XTBG, secondary tropical montane evergreen broad-leaved forest, 21°57.534'N, 101°12.300'E, ca. 860 m, 4.VIII.2007, G. Zheng leg.

Diagnosis and description

See Liu et al. (2016). Male palp as in Figs 7, 56B, 66B, epigyne as in Figs 8A–D, 75D, 83D, 91D, habitus as in Fig. 8E–H.

Distribution

China (Yunnan).

Most similar species

Clubiona submoralis.

Clubiona parconcinna Deeleman-Reinhold, 2001

Figs 9, 10, 55C, 65C, 75C, 83C, 91C

Clubiona parconcinna Deeleman-Reinhold, 2001: 117, figs 34–40 (♂♀); Dankittipakul and Singtripop 2008b: 645, figs 1–9 (♂).

Material examined

1♂ (YHCLU0143), China, Yunnan Province: Xishuangbanna: Mengla County: Bubang Village: monsoon forest, 21°36.827'N, 101°34.847'E, ca. 690 m, 12.VIII.2012, G. Zheng et al. leg; 1♀ (YHCLU0260), Mengla County: Manda Village: secondary forest, 22°01.421'N, 101°23.418'E, ca. 1188 m, 28.VII.2012, Q.Y. Zhao and Z.G. Chen leg; 6♂7♀, Menglun Town: XTBG, 55th km landmark in the Menglun Nature Reserve, tropical ravine rainforest, 21°54.883'N, 101°12.147'E, ca. 829 m, 15.VIII.2011, Q.Y. Zhao and Z.G. Chen leg.

Diagnosis and description

See Deeleman-Reinhold (2001) and Dankittipakul and Singtripop (2008b). Male palp as in Figs 9, 55C, 65C, epigyne as in Figs 10A–D, 75C, 83C, 91C, habitus as in Fig. 10E–H.

Distribution

Thailand (Nakhon Ratchasima), Indonesia (Borneo), China (Yunnan).

Most similar species

Clubiona xiaoci sp. nov.

Clubiona pollicaris Wu, Zheng & Zhang, 2015

Figs 11, 12, 56C, 66C, 76B, 84B, 92B

Clubiona pollicaris Wu et al., 2015: 20, figs 13–19, 23–27 (♂♀).

Material examined

1♂ (YHCLU0020), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Rainforest Nature Park, 21°55.017'N, 101°16.450'E, ca. 572 m, 16.VII.2018, H. Yu leg; 1♀ (YHCLU0021), XTBG, Paramichelia baillonii forest, 21°54.772'N, 101°16.043'E, ca. 556 m, 19.VII.2018, H. Yu leg; 16♂32♀, XTBG, low evergreen forest, 21°53.794'N, 101°17.152'E, ca. 594 m, 27.XI.2009, G. Tang and Z.Y. Yao leg.

Diagnosis and description

See Wu et al. (2015). Male palp as in Figs 11, 56C, 66C, epigyne as in Figs 12A–D, 76B, 84B, 92B, habitus as in Fig. 12E–H.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona globosa.

Clubiona rama Dankittipakul & Singtripop, 2008

Figs 13, 14, 53B, 63B

Clubiona rama Dankittipakul & Singtripop, 2008b: 645, figs 10–23 (♂♀); Dhali et al. 2016: 289, figs 7A–E, 8A–C (♀); Dhali et al. 2017: 58, figs 237–241, pl. 21 (♀); Yu et al. 2017b: 692, figs 6–10 (♀).

Material examined

1♂, China: Yunnan Province: Xishuangbanna: Jinghong City: Nabanhe Natural Reserve, Mandian Waterfall, monsoon forest, 22°7.845'N, 100°39.749'E, ca. 736 m, 22.VIII.2012, G. Zheng leg.

Diagnosis and description

See Dankittipakul and Singtripop (2008b). Male palp as in Figs 13, 53B, 63B, habitus as in Fig. 14.

Distribution

India (West Bengal), Thailand (Phitsanulok), China (Yunnan).

Most similar species

Clubiona subrama.

Clubiona subdidentata Yu & Li, sp. nov.

Figs 15, 74F, 82F, 90F

Holotype

♀ (IZCAS-Ar 34749), China: Yunnan Province: Xishuangbanna: Mengla County: Xiaolongha Village: 22°5.017'N, 100°22.084'E, ca. 1118 m, 24.VII.2012, Q.Y. Zhao and Z.G. Chen leg. Other material examined. 1♀ (YHCLU0073), same data as holotype

Etymology

The specific name is taken from its similarity to C. didentata; adjective.

Diagnosis

The female of C. subdidentata sp. nov. can be distinguished from all other members of the C. corticalis group with the exception of C. didentata (Yu and Li 2019b: 207, figs 6A–D; Figs 74E, 82E, 90E) by having an atrial membrane (atrial membrane is absent in almost all species of the corticalis group) and similar vulva but can be recognised by the nearly equilateral triangular atrial membrane (Figs 15A–C, 74F, 82F) (vs. tongue- shaped in C. didentata; Figs 74E, 82E) and by the distinct copulatory ducts (Figs 15D, E, 90F) (copulatory ducts absent in C. didentata; Fig. 90E).

Description

Female. Holotype (Fig. 15F, G): Total length 3.63; carapace 1.45 long, 1.14 wide; opisthosoma 2.18 long, 1.32 wide. Carapace uniformly greyish white, without any pattern or markings; ocular region distinctly narrowed, cervical groove and radial grooves indistinct; tegument smooth, all setae detached in ethanol. Eyes: AER slightly recurved, PER wider than AER and almost straight in dorsal view. Eye sizes and interdistances: AME 0.09, ALE 0.09, PME 0.09, PLE 0.07, AMEAME 0.06, AMEALE 0.07, PMEPME 0.19, PMEPLE 0.10, MOQL 0.21, MOQA 0.23, MOQP 0.38. Chelicerae light orange, with three promarginal and two retromarginal teeth. Sternum pale brown, 0.80 long, 0.65 wide. Labium and endites coloured as chelicerae. Legs greyish white, uniformly coloured. Leg measurements: I 2.73 (0.81, 1.15, 0.55, 0.23), II 3.15 (0.93, 1.25, 0.74, 0.22), III 2.85 (0.93, 0.95, 0.70, 0.29), IV 3.39 (1.19, 1.32, 1.05, 0.39). Abdomen oval, nearly pure white, with inconspicuous anterior setal tufts, dorsum with two pairs of inconspicuous muscle depressions; venter without pattern.

Epigyne (Figs 15A–E, 74F, 82F, 90F). Epigynal plate ca. 1.5 × wider than long, margin not delimited; spermathecae and bursae prominently visible through epigynal plate. Atrium small, anteriorly covered by an atrial membrane. Atrial membrane shaped nearly like an equilateral triangle, with a blunt apex. Copulatory openings small, located at basolateral atrial borders. Copulatory ducts distinct, extend transversally, connecting to posteriorly located bursae. Spermathecae small, consisting of a bean-shaped proximal part and an acicular distal part, with short fertilisation ducts terminally. Bursae reniform, close together, ca. 1.3 × longer than wide, bursal surface hyaline and smooth, inside pigmented and sclerotised.

Male. Unknown.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona submoralis Wu, Zheng & Zhang, 2015

Figs 16, 17, 55B, 65B, 75B, 83B, 91B

Clubiona submoralis Wu et al., 2015: 17, figs 1–12 (♂♀).

Material examined

1♂ (YHCLU0028), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Paramichelia baillonii Forest, 21°54.772'N, 101°16.043'E, ca. 556 m, 19.VII.2018, H. Yu leg; 1♀ (YHCLU0029), XTBG, rubber plantation, 21°54.674'N, 101°16.207'E, ca. 583 m, 21.VII.2018, H. Yu leg.; 6♂8♀, XTBG, secondary forest, 21°54.459'N, 101°16.755'E, ca. 644 m, 20.XI.2009, G. Tang and Z.Y. Yao leg.

Diagnosis and description

See Wu et al. (2015). Male palp as in Figs 16, 55B, 65B, epigyne as in Figs 17A–D, 75B, 83B, 91B, habitus as in Fig. 17E–H.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona moralis.

Clubiona subrama Yu & Li, 2019

Figs 53C, 63C, 73E, 81E, 89E

Clubiona subrama Yu & Li, 2019a: 153, figs 3A–E, 4A–G (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34524), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Lvshilin Forest Park, limestone tropical seasonal rainforest, 21°54.705'N, 101°16.898'E, ca. 656 m, 13.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♀, XTBG, rubber plantation, 21°54.498'N, 101°16.326'E, ca. 586 m, 29.II.2009, G. Zheng leg; 1♂ (YHCLU0083) and 1♀ (YHCLU0084), XTBG, Lvshilin Forest Park, evergreen forest, 21°54.555'N, 101°16.860'E, ca. 616 m, 29.XI.2009, G. Tang and Z.Y. Yao leg.

Diagnosis and description

See Yu and Li (2019a). Male palp as in Figs 53C, 63C, epigyne as in Figs 73E, 81E, 89E.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona rama.

Clubiona subyaginumai Yu & Li, 2019

Figs 54C, 64C, 75F, 83F, 91F

Clubiona subyaginumai Yu & Li, 2019a: 158, figs. 5A–E, 6A–G (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34548), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary tropical forest, 21°54.380'N, 101°16.815'E, ca. 627 m, 23.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♀, XTBG, bamboo plantation, 21°54.380'N, 101°16.815'E, ca. 620 m, 21.XI.2009, G. Tang and Z.Y. Yao leg.

Diagnosis and description

See Yu and Li (2019a). Male palp as in Figs 54C, 64C, epigyne as in Figs 75F, 83F, 91F.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona yaginumai.

Clubiona tixing Yu & Li, sp. nov.

Figs 18, 73D, 81D, 89D

Holotype

♀ (IZCAS-Ar 34750), China: Yunnan Province: Xishuangbanna: Jinghong City: Nabanhe Natural Reserve, Mandian Waterfall, monsoon forest, 22°7.845'N, 100°39.749'E, ca. 736 m, 22.VIII.2012, G. Zheng leg.

Etymology

The specific name is derived from the Chinese pinyin tī xíng, which means ‘trapezoid’, referring to the trapezoidal atrium; noun in apposition.

Diagnosis

Females of this species resemble those of C. zhangmuensis (Zhang et al. 2007: 38, figs 7–8) in having a similar atrium and endogyne but can be recognised by the presence of an atrial membrane (Figs 18A–C, 73D, 81D) (vs. absent; Zhang et al. 2007: 38, fig. 7).

Description

Female. Holotype (Fig. 18F, G): Total length 6.62; carapace 2.80 long, 2.02 wide; opisthosoma 3.82 long, 2.28 wide. Carapace brown in alcohol, uniformly coloured, without distinct pattern, ocular region distinctly narrowed, cervical groove and radial grooves indistinct; tegument smooth, clothed with short setae. Eyes: in dorsal view, both AER and PER almost straight, the former narrower than the latter. Eye sizes and interdistances: AME 0.14, ALE 0.15, PME 0.14, PLE 0.12, AMEAME 0.12, AMEALE 0.13, PMEPME 0.39, PMEPLE 0.24, MOQL 0.43, MOQA 0.41, MOQP 0.67. Chelicerae robust and dark brown, with five promarginal and three retromarginal teeth. Sternum yellowish brown, darker marginally, 1.63 long, 1.00 wide. Labium and endites coloured as carapace. Legs uniformly yellowish white. Leg measurements: I – (1.71, 2.25, –, –), II – (1.78, 2.42, –, –), III – (1.60, 1.88, –, –), IV – (2.11, 2.55, 1.17, –). Abdomen elongate-oval, uniformly pale brown, with numerous brown spots; dorsally with a lengthwise white heart mark, reaching posterior half, with two pairs of muscle depressions located at lateral part of heart mark; venter medially with four longitudinal dotted lines.

Epigyne (Figs 18A–E, 73D, 81D, 89D). Epigynal plate distinctly wider than long, margin not rebordered, spermathecae and bursae are indistinctly visible through epigynal plate in ventral view. Atrium large, more or less rectangular or trapezoidal, ca. 1.3 × wider than long, ca. 1/3 of epigyne length, with tongue-shaped membrane on anterior margin, posterior atrial border not delimited. Copulatory openings indistinct, located at anterolateral atrial borders. Copulatory ducts short, bent dorsally and ascend obliquely to connect with bursae. Spermathecae convoluted, anteriorly situated, spermathecal heads thick. Bursae reniform, close together, ca. 1.6 × longer than wide, surface translucent and wrinkled.

Male. Unknown.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona tiane Yu & Li, 2019

Figs 19, 54D, 64D, 74D, 82D, 90D

Clubiona tiane Yu & Li, 2019b: 204, figs 3A–E, 4A–C (♂).

Material examined

Types. Holotype ♂ (IZCAS Ar 34703), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, bamboo plantation, 21°53.642'N, 101°16.940'E, ca. 589 m, 22.VII.2018, H. Yu and Z.G. Chen leg. Other material examined. 1♂ (YHCLU0054), XTBG, rubber-tea plantation, 21°55.233'N, 101°15.850'E, ca. 572 m, 16.VII.2018, Hao Yu et al. leg; 1♀ (YHCLU0053), XTBG, Rainforest Nature Park, 21°55.017'N, 101°16.450'E, ca. 572 m, 16.VII.2018, H. Yu et al. leg.

Diagnosis

Females of C. tiane resemble those of C. dactylina (Liu et al. 2016: figs 18, 19, 23, 24) in having a semi-circular atrium and tubular spermathecae but differ in the following: (1) indistinct copulatory openings hidden at anterior sides of atrial border (Figs 19A–C, 74D, 82D) (vs. distinct copulatory openings located posteriorly in atrium; Liu et al. 2016: figs 18, 23); (2) more or less lengthwise, thick spermathecae (Figs 19D, E, 90D) (vs. nearly horizontal and relatively thin spermathecae); (3) distinct and thick spermathecal heads (Figs 19D, E, 90D) (vs. indistinct and relatively small spermathecal heads; Liu et al. 2016: figs 19, 24). Males of C. tiane can be easily recognised by the curved embolus shaped like a swan’s neck (Figs 54D, 64D).

Description

Male. See Yu and Li (2019b). Male palp as in Figs 54D, 64D.

Female (Fig. 19F, G). Total length 3.90; carapace 1.77 long, 1.06 wide; opisthosoma 2.14 long, 0.93 wide. Carapace light grey except brownish ocular area, without pattern; ocular region slightly narrowed, cervical groove and radial grooves indistinguishable; tegument smooth, all setae detached in ethanol. Eyes: both AER and PER slightly recurved in dorsal view, PER slightly wider than AER. Eye sizes and interdistances: AME 0.09, ALE 0.09, PME 0.10, PLE 0.08, AMEAME 0.09, AMEALE 0.06, PMEPME 0.17, PMEPLE 0.12, MOQL 0.23, MOQA 0.31, MOQP 0.32. Chelicerae coloured as ocular area, with six promarginal and four retromarginal teeth. Sternum yellowish white, 0.94 long, 0.61 wide. Labium and endites brownish. Legs yellowish white, without distinct markings. Leg measurements: I 2.38 (0.68, 1.12, 0.28, 0.30), II 2.87 (0.87, 1.25, 0.35, 0.39), III 2.30 (0.63, 0.63, 0.79, 0.25), IV 3.78 (1.39, 1.19, 0.88, 0.32). Abdomen elongate, oval, nearly pure white, with inconspicuous anterior setal tufts, dorsum clothed with dense setae; venter uniformly white, without pattern.

Epigyne (Figs 19A–E, 74D, 82D, 90D). Epigynal plate slightly wider than long, margin not rebordered, spermathecae and bursae obscured through epigynal plate in ventral view. Atrium small, semi-circular, ca. 1/4 of epigyne width, anterior atrial border heavily sclerotised. Copulatory openings indistinct, hidden by anterior margin of atrium. Copulatory ducts indistinct, directed posteriorly to connect with spermathecae. Spermathecae longitudinal, with short, acicular fertilisation ducts. Spermathecal heads digitiform and horizontal, arising on the proximal part of the spermathecae. Bursae ovoid, large, close together, ca. 1.7 × longer than wide, surface translucent, with a wrinkled and ribbed appearance.

Distribution

Known only from Xishuangbanna.

Remarks

The female of the species is described for the first time.

Clubiona xiaoci Yu & Li, sp. nov.

Figs 20, 21, 55D, 65D, 75E, 83E, 91E

Holotype

♂ (IZCAS-Ar 34751), China: Yunnan Province: Xishuangbanna: Jinghong City: Menga Town: Wengnan Village: secondary forest, 22°5.020'N, 100°22.087'E, ca. 1118 m, 24.VII.2012, Q.Y. Zhao and Z.G. Chen leg. Paratype: 1♀ (IZCAS-Ar 34752), same data as holotype. Other material examined. 1 ♂ (YHCLU0088) and 1 ♀ (YHCLU0089), same locality and same collectors as holotype, 22°4.997'N, 100°22.223'E, ca. 1137 m, 25.VII.2012.

Etymology

The specific name is derived from the Chinese pinyin xiǎo cì, which means ‘small spines’, referring to the short spines located on the palpal tibia and patella; noun in apposition.

Diagnosis

Clubiona xiaoci sp. nov. is very similar to C. parconcinna (see Figs 9, 10, 55C, 65C, 75C, 83C, 91C and Deeleman-Reinhold 2001: 117, figs 34–40). Males are similar by the palpal tibia with several short spines. Females of C. xiaoci sp. nov. resemble those of C. parconcinna in having similar atrial anterior margins and globular spermathecae. C. xiaoci sp. nov. can be distinguished from C. parconcinna by the following characters: for the males, conductor absent in C. xiaoci sp. nov. (Figs 20, 55D, 65D) (vs. distinct; Figs 9B, D, E, 55C, 65C), retrolateral patellar apophysis with short, modified spines in new species (Figs 20B, 55D, 65D) (vs. without spines; Figs 9B, 55C, 65C); for the females, copulatory openings are partly fused in C. xiaoci sp. nov. (Figs 21A, B, 75E, 83E) (vs. copulatory openings separated; Figs 10A, B, 75C, 83C), copulatory ducts distinctly shorter and thicker in C. xiaoci sp. nov. (cf. Figs 21C, D, 91E and Figs 10C, D, 91C).

Description

Male. Holotype (Fig. 21E, F): Total length 3.17; carapace 1.51 long, 1.10 wide; opisthosoma 1.66 long, 0.94 wide. Carapace greyish white, slightly lighter in cephalic area, with a pair of short faint lines running longitudinally from behind AME, ocular region distinctly narrowed; cervical groove indistinct; tegument smooth, clothed with short setae. Eyes: AER slightly recurved, PER almost straight, the latter wider than the former. Eye sizes and interdistances: AME 0.07, ALE 0.08, PME 0.07, PLE 0.06, AMEAME 0.07, AMEALE 0.05, PMEPME 0.10, PMEPLE 0.07, MOQL 0.19, MOQA 0.23, MOQP 0.31. Chelicerae robust and light orange, with three promarginal and two retromarginal teeth. Sternum pale yellow, 0.83 long, 0.63 wide. Labium and endites coloured as chelicerae. Legs yellowish white, without markings. Leg measurements: I – (1.06, 1.41, 0.75, –), II 4.22 (1.19, 1.76, 0.83, 0.44), III 3.02 (0.93, 1.12, 0.75, 0.22), 4.44 (1.27, 1.43, 1.27, 0.46). Abdomen oval, dorsally light pink with conspicuous anterior setal tufts; venter pale yellow, without pattern.

Palp (Figs 20A–E, 55D, 65D). Femur unmodified. Patella with single retrolateral apophysis, apophysis with short, modified spines at apex. Tibia short, cup-shaped, retrolaterally with several short spines near base, with two apophyses: a papilliform, partly membranous ventro-retrolateral apophysis and a dorsal one, with a blunt apex, trapezoidal. Cymbium 2.3 × longer than wide. Tegulum elongate, oval, and bulging, 1.5 × longer than wide; sperm duct indistinct in ventral view. Embolus needle-like, distinctly short, originating at distal portion of tegulum, gradually tapering toward tip, apex sharp and prolaterally pointed; embolar base represented by enlarged tubercle.

Female. Paratype (Fig. 21G, H): total length 3.93; carapace 1.69 long, 1.21 wide; opisthosoma 2.24 long, 1.30 wide. Eye sizes and interdistances: AME 0.09, ALE 0.10, PME 0.10, PLE 0.09, AMEAME 0.05, AMEALE 0.05, PMEPME 0.19, PMEPLE 0.09, MOQL 0.28, MOQA 0.23, MOQP 0.38. Sternum 0.99 long, 0.67 wide. Leg measurements: I 3.56 (1.11, 1.42, 0.63, 0.41), II 3.80 (1.15, 1.52, 0.67, 0.44), III – (0.90, –, 0.78, 0.34), IV 4.42 (1.32, 1.48, 1.15, 0.47). Distinctly larger and darker than male, other characters as in male.

Epigyne (Figs 21A–D, 75E, 83E, 91E). Epigynal plate nearly as wide as long, spermathecae distinctly visible through integument. Atrium indistinct, anterior margin rebordered. Copulatory openings small and partly fused, situated anteriorly on atrium. Copulatory ducts short, ascending obliquely, connecting to posteriorly located bursae then ascending to anteriorly located spermathecae. Spermathecae nearly globular, separated by 1.5 diameters. Fertilisation ducts small, acicular, and membranous, located on posterior surface of spermathecae. Bursae reniform, large, close together, ca. 1.25 × longer than wide, surface translucent and wrinkled.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona xiaokong Yu & Li, sp. nov.

Figs 22, 74A, 82A, 90A

Holotype

♀ (IZCAS-Ar 34753), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Anogeissus acuminata plantation (ca. 20 yr.), 21°53.819'N, 101°17.075'E, ca. 609 m, 27.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♀ (YHCLU0078), XTBG, secondary tropical forest, 21°54.833'N, 101°16.781'E, ca. 617 m, 26.IV.2019, Z.G. Chen leg.

Etymology

The specific name is derived from the Chinese pinyin xiǎo kǒng, which means small opening, referring to the small atrium; noun in apposition.

Diagnosis

The new species is similar to C. falciforma (Liu et al. 2016: 567, figs 30, 31, 35, 36) by the ɔc-shaped spermathecae. From C. falciforma, the female of the new species can be easily distinguished by: the nearly trapezoidal atrium distinctly longer than wide (Figs 22A–C, 74A, 82A) (vs. elliptical atrium wider than long; Liu et al. 2016: figs 30, 35); copulatory openings located at lateral atrial borders (Figs 22A–C, 74A, 82A) (vs. situated basolaterally in atrium; Liu et al. 2016: figs 30, 35); (and the transverse copulatory ducts (Figs 22D, E, 90A) (vs. copulatory ducts descending longitudinally; Liu et al. 2016: figs 31, 36).

Description

Female. Holotype (Fig. 22F, G): Total length 6.48; carapace 3.28 long, 2.26 wide; opisthosoma 3.19 long, 2.03 wide. Carapace elongate, oval, light brown, uniformly coloured, without pattern, fovea red; pars cephalica slightly narrowed, cervical groove indistinct; tegument smooth, with erect, thin, dark setae on front ridge. Eyes: In dorsal view, AER slightly recurved, PER almost straight, PER wider than AER. Eye sizes and interdistances: AME 0.13, ALE 0.16, PME 0.14, PLE 0.12, AMEAME 0.16, AMEALE 0.11, PMEPME 0.35, PMEPLE 0.25, MOQL 0.36, MOQA 0.46, MOQP 0.68. Chelicerae protruding and robust, coloured as carapace, with long, orange fangs, with five promarginal and two retromarginal teeth. Sternum nearly white, 1.60 long, 1.10 wide. Labium and endites coloured as carapace. Legs light coloured, dorsally slightly darker, without markings. Leg measurements: I 6.23 (1.76, 2.60, 1.18, 0.68), II 6.87 (2.04, 2.68, 1.34, 0.81), III 5.97 (1.85, 2.15, 1.39, 0.57), IV 8.26 (2.35, 2.72, 2.41, 0.78). Abdomen oval and light brown, without pattern, dorsum densely covered with long, dark setae on a light background; venter sparsely covered with short, white setae.

Epigyne (Figs 22A–E, 74A, 82A, 90A). Epigynal plate slightly longer than wide, margin not rebordered; spermathecae and bursae indistinctly visible through transparent integument. Atrium represented by small pore, ca. 2 × longer than wide, ca. 1/6 of epigyne length and 1/11 of epigyne width. Copulatory openings large, situated laterally on the atrium. Copulatory ducts short and thick, heavily sclerotised, expanding laterally, then connecting to tubular spermathecae. Spermathecae long, consisting of smooth proximal half and wrinkled distal half, with small fertilisation ducts terminally, proximal half shaped like ɔc, the distal half irregularly shaped. Bursae reniform, close together, distinctly larger than spermathecae, 1.3 × longer than wide, surface translucent and smooth.

Male. Unknown.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona yejiei Yu & Li, sp. nov.

Figs 23, 73C, 81C, 89C

Holotype

♀ (IZCAS-Ar 34754), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Paramichelia baillonii plantation, 21°53.823'N, 101°17.072'E, ca. 613 m, 1–17.II.2007, G. Zheng leg.

Etymology

This species is named after Mr. Yejie Lin (Beijing City, China) who has helped us greatly with this research.

Diagnosis

The new species is similar to C. cochlearis (Figs 1A–E, 73A, 81A, 89A) in the general appearance of the atrium and endogyne. From C. cochlearis, the female of the new species can be easily distinguished by the shape of the atrium and spermathecae: (1) atrium ellipsoid (Figs 23A–C, 73C, 81C) (vs. atrium apple-shaped; Figs 1A–C, 73A, 81A); (2) spermathecae thicker and not twisted, diameter ca. 1/3 of atrium length (Figs 23D, E, 89C) (vs. thinner and strongly convoluted, diameter not more than 1/8 of atrium length; Figs 1D, E, 89A).

Description

Female. Holotype (Fig. 23F, G). Total length 9.47; carapace 4.47 long, 2.56 wide; opisthosoma 5.50 long, 3.82 wide. Carapace brownish red, pars cephalica slightly darker in ocular area, without markings, cephalic region slightly narrowed, cervical groove indistinct; tegument smooth, covered with numerous fine setae. Eyes: AER slightly recurved, PER slightly procurved, PER wider than AER. Eye sizes and interdistances: AME 0.17, ALE 0.17, PME 0.17, PLE 0.09, AMEAME 0.25, AMEALE 0.85, PMEPME 0.45, PMEPLE 0.47, MOQL 0.55, MOQA 0.58, MOQP 0.79. Chelicerae robust and protruding, coloured as ocular area, with four promarginal and two retromarginal teeth. Sternum centrally orange and marginally red, 2.13 long, 1.36 wide. Labium and endites coloured as chelicerae. Legs light orange, uniformly coloured, without pattern. Leg measurements: I 8.64 (2.33, 3.63, 2.03, 0.64), II 8.73 (2.76, 3.60, 1.44, 0.93), III 8.16 (2.51, 2.96, 1.77, 0.92), IV 10.54 (3.32, 3.48, 2.60, 1.14). Abdomen uniformly cream coloured, dorsum with two pairs of conspicuous muscle depressions; venter medially with two longitudinal broken lines.

Epigyne (Figs 23A–E, 73C, 81C, 89C). Epigynal plate slightly longer than wide, through which bursae and copulatory ducts are easily visible. Atrium shaped like an irregular ellipse, ca. 1.3 × wider than long, ca. 1/5 epigyne length. Copulatory openings indistinct, located at basolateral atrial borders. Copulatory ducts short and thick, heavily sclerotised, running laterally, bent 90 degrees dorsally and then obliquely descending, connecting with tubular spermathecae. Spermathecae long and sinuous, with even thickness throughout. Fertilisation ducts acicular, membranous, located terminally on spermathecae. Bursae large and oblong, close together, ca. 1.5 × longer than wide, surface translucent and wrinkled.

Male. Unknown.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona zhaoi Yu & Li, sp. nov.

Figs 24, 74B, 82B, 90B

Holotype

♀ (IZCAS-Ar 34755), China, Yunnan Province: Xishuangbanna: Mengla County: Xiaolongha Village: biodiversity preservation corridor, 21°24.798'N, 101°37.880'E, ca. 693 m, 28.VI.2012, Q.Y. Zhao and C.X. Gao leg. Other material examined. 1♀ (YHCLU0086), same locality, same time, and same collectors as holotype, 21°24.408'N, 101°37.827'E, ca. 662 m.

Etymology

The specific name is a patronym after Qingyuan Zhao (Beijing City, China), collector of several specimens examined in this study.

Diagnosis

Females of the new species are easily distinguished from others in the species group, with the exception for C. altissimoides (Liu et al. 2007: 65, figs 6, 7), by the similarly shaped atria and the general shape of the endogyne. Clubiona zhaoi sp. nov. can be separated from C. altissimoides by the copulatory openings located at the anterior atrial margins (Figs 24A–C, 74B, 82B) (vs. located at the underside of the atrium; Liu et al. 2007: fig. 6) and by having the spermathecae distinctly smaller than the bursae (Figs 24D, E, 90B) (vs. larger; Liu et al. 2007: fig. 7).

Description

Female. Holotype (Fig. 24F, G): Total length 5.82; carapace 2.80 long, 1.98 wide; opisthosoma 3.02 long, 1.65 wide. Carapace light brown except dark brown ocular area, without distinct pattern; fovea dark; cephalic region distinctly narrowed, cervical groove distinct, radial grooves inconspicuous; tegument smooth, clothed with short, fine setae. Eyes: AER slightly recurved, PER slightly wider than AER, almost straight in dorsal view. Eye sizes and interdistances: AME 0.11, ALE 0.15, PME 0.12, PLE 0.08, AMEAME 0.11, AMEALE 0.08, PMEPME 0.32, PMEPLE 0.19, MOQL 0.39, MOQA 0.37, MOQP 0.59. Chelicerae robust and brownish red, both margins with four teeth. Sternum yellowish brown, 1.38 long, 0.95 wide. Labium and endites coloured as carapace. Legs yellowish, without distinct markings. Leg measurements: I – (1.58, –, –, –), II 6.60 (1.89, 2.52, 1.38, 0.81), III 5.29 (1.64, 1.69, 1.43, 0.53), IV – (–, –, 1.17, –). Abdomen long, oval, dorsally grey with dense setae and a lengthwise white heart mark, reaching posterior half; with a pair of muscle depressions located at distal part of heart mark; venter off-white.

Epigyne (Figs 24A–E, 74B, 82B, 90B). Epigynal plate nearly as broad as long, spermathecae and bursae indistinctly visible through integument. Atrium small, ca. 1/4 of epigyne width, with M-shaped anterior margin (or hood), without posterior margin. Copulatory openings small but distinct, circular, located on anterior part of atrium. Copulatory ducts indistinct. Spermathecae consisting of fan-shaped proximal part and convoluted distal part, with small fertilisation ducts terminally; the two spermathecae separated by 0.8 diameters. Bursae oval, close together, ca. 1.4 × longer than wide, surface translucent and wrinkled.

Male. Unknown.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona zhigangi Yu & Li, sp. nov.

Figs 25, 26, 53D, 63D, 73F, 81F, 89F

Holotype

♂ (IZCAS-Ar 34756, YHCLU0185), China: Yunnan Province: Xishuangbanna: Mengla County: Menga Town: XTBG, secondary tropical forest, 21°53.634'N, 101°17.172'E, ca. 620 m, 28. IV.2019, Z.G. Chen et al. leg. Paratype: 1♀ (IZCAS-Ar 34757), XTBG, secondary tropical seasonal moist forest, 21°54.718'N, 101°16.940'E, ca. 645 m, 27. VII.2007, G. Zheng leg. Other material examined. 1♀ (YHCLU0138), China: Yunnan Province: Xishuangbanna: Mengla County: Menga Town: XTBG, Paramichelia baillonii plantation, 21°53.823'N, 101°17.072'E, ca. 613 m, 15. IV.2007, G. Zheng leg.

Etymology

The specific name is a patronym after Zhigang Chen (Beijing City, China), collector of several specimens used in this study.

Diagnosis

Males of C. zhigangi sp. nov. resemble those of C. subrama (Yu and Li 2019a: 153, fig. 3A–E; Figs 53C, 63C) in having similar retrolateral and ventral tibial apophyses and a slender embolus but differ by: (1) a thicker embolus, with a sinuate apex (Figs 25A–E, 53D, 63D) (vs. filiform embolus, thinner, embolar tip not curved; Figs 53C, 63C); (2) short conductor, ca. 1/5 of tegulum length, with a blunt tip (Fig. 25C–E) (vs. conductor long, not less than 1/3 of tegulum length, with a sharp tip; Fig. 53C). Females also resemble those of C. subrama in having an anteriorly cordiform and posteriorly elongate, narrowed atrium and the general shape of the endogyne but can be distinguished from the latter by the distinctly shorter copulatory ducts not convoluted (Figs 26C, D, 89F) (vs. long copulatory ducts strongly entwined, moving longitudinally and expanding obliquely 2 ×, respectively, forming two horizontal loops; Fig. 89E).

Description

Male. Holotype (Fig. 26E, F). Total length 5.91; carapace 2.85 long, 2.17 wide; opisthosoma 3.06 long, 1.88 wide. Carapace brown, darker in the front, without distinct pattern, fovea red; cephalic region distinctly narrowed, cervical groove and radial grooves indistinct; tegument smooth, clothed with short, fine setae. Eyes: AER slightly recurved, PER wider than AER and slightly procurved in dorsal view. Eye sizes and interdistances: AME 0.13, ALE 0.12, PME 0.14, PLE 0.11, AMEAME 0.19, AMEALE 0.13, PMEPME 0.30, PMEPLE 0.24, MOQL 0.36, MOQA 0.32, MOQP 0.53. Chelicerae robust, red wine coloured, with three promarginal and two retromarginal teeth. Sternum pale brown, 1.64 long, 1.02 wide. Labium and endites coloured as carapace. All legs missing. Abdomen elongate, oval, dorsal scutum trapezoidal, lightly sclerotised, with a thick tuft of setae anteriorly; dorsum brown, with dense setae, with a pair of muscle depressions located at central part of dorsal scutum; venter grey.

Palp (Figs 25A–E, 53D, 63D). Femur and patella unmodified. Tibia short, ca. 1/2 of cymbium length, with 2 apophyses: a large, semi-circular ventral one, ca. 1/3 of palpal tibia length, and a relatively small, claw-shaped retrolateral apophysis. Tegulum elongate, oval, 1.9 × longer than wide; bulb strongly bulging and prolapsed, sperm duct indistinct in ventral view. Embolus slender, originating from prolateral side of tegulum, ca. 1/2 of tegulum length, tip sinuate and extending above the apex of the cymbium. Conductor short, membranous, ca. 1/3 of the embolus length, originating from retrolateral side of tegulum, with basal torsion and a distal finger-like point, tip hidden behind embolus.

Female. Paratype (Fig. 26G, H). Total length 8.51; carapace 3.56 long, 2.48 wide; opisthosoma 4.95 long, 3.05 wide. Eye sizes and interdistances: AME 0.16, ALE 0.19, PME 0.15, PLE 0.15, AMEAME 0.12, AMEALE 0.11, PMEPME 0.35, PMEPLE 0.28, MOQL 0.36, MOQA 0.46, MOQP 0.65. Sternum 1.88 long, 1.14 wide. Leg measurements: I 5.99 (1.17, 2.33, 1.13, 0.74), II 6.74 (1.99, 2.67, 1.29, 0.79), III 5.89 (1.74, 2.11, 1.51, 0.54), IV 8.05 (2.31, 2.82, 2.30, 0.62). Distinctly larger and darker than male, other characters as in male.

Epigyne (Figs 26A–E, 73F, 81F, 89F). Epigynal plate slightly wider than long, spermathecae indistinctly visible and bursae prominently visible through integument in ventral view. Atrium small and elongate, narrow, ca. 1/2 × epigyne length and 1/7 × epigyne width, atrial anterior margin M-shaped, posterior margin not delimited. Copulatory openings small, located on anterolateral margin of atrium. Copulatory ducts directed laterally then ascending obliquely, connecting to spermathecae at central axis of the vulva. Spermathecae tubular, long and sinuous, strongly twisted. Fertilisation ducts acicular, curved, located terminally on spermathecae. Bursae reniform, large, close together, ca. 1.5 × longer than wide, surface translucent and wrinkled.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona ternatensis group

Hirtia Thorell, 1881: 222 (type species H. ternatensis Thorell, 1881).

Clubiona: Simon 1897: 76 (synonymised Hirtia).

Clubiona hystrix group: Deeleman-Reinhold 2001: 90.

Diagnosis

See Deeleman-Reinhold (2001) and Yu and Li (2019a).

Description

See Deeleman-Reinhold (2001).

Composition and distribution

The Clubiona ternatensis group tentatively contains 33 species (see Table 4).

Comments

This group was defined by Deeleman-Reinhold (2001) for Oriental species. However, the three first described species were placed in the genus Hirtia Thorell, 1881, and H. ternatensis was chosen as the type species by Thorell (1881). Simon (1897) synonymised Hirtia with Clubiona. The recently discovered species of the group are strikingly different from Clubiona pallidula (Clerck, 1757) (type species of Clubiona). It seems that Hirtia should be resurrected in the future.

Table 4.

Species in the Clubiona ternatensis group.

Species name Known sex Distribution
1 C. analis Thorell, 1895 India (West Bengal), Bangladesh (Barisal), Myanmar (Moulmein)
2 C. bachmaensis Ono, 2009 Vietnam
3 C. bengalensis Biswas, 1984 India (Maharashtra)
4 C. brevispina Huang & Chen, 2012 ♂♀ China (Taiwan)
5 C. chathamensis Simon, 1905 New Zealand (Chatham Is.)
6 C. damirkovaci Deeleman-Reinhold, 2001 ♂♀ Malaysia (Kuala Lumpur)
7 C. ericius Chrysanthus, 1967 ♂♀ New Guinea
8 C. esuriens Thorell, 1897 Myanmar (specific locality not clear)
9 C. hatamensis (Thorell, 1881) New Guinea
10 C. heteroducta Zhang & Yin, 1998 China (Yunnan)
11 C. hitchinsi Saaristo, 2002 ♂♀ Seychelles, French Polynesia
12 C. hystrix Berland, 1938 ♂♀ Indonesia (Lesser Sunda Is.)
13 C. jaegeri Ono, 2011 Palau Is.
14 C. kapataganensis Barrion & Litsinger, 1995 Philippines (Laguna)
15 C. kowong Chrysanthus, 1967 ♂♀ New Guinea
16 C. kuu Jäger & Dankittipakul, 2010 Laos
17 C. maipai Jäger & Dankittipakul, 2010 ♂♀ Thailand (Mae Hong Son)
18 C. meraukensis Chrysanthus, 1967 ♂♀ New Guinea
19 C. oceanica Ono, 2011 ♂♀ Japan (Chichijima Is.)
20 C. pantherina Chrysanthus, 1967 ♂♀ New Guinea
21 C. papuana Chrysanthus, 1967 New Guinea
22 C. paranghinlalakirta Barrion & Litsinger, 1995 Philippines (Misamis Oriental)
23 C. pseudomaxillata Hogg, 1915 New Guinea
24 C. pseudopteroneta Raven & Stumkat, 2002 ♂♀ Australia (Queensland)
25 C. ramoiensis (Thorell, 1881) New Guinea
26 C. sertungensis Hayashi, 1996 ♂♀ Indonesia (Krakatau)
27 C. subkuu Yu & Li, 2019 ♂♀ China (Yunnan)
28 C. ternatensis (Thorell, 1881) Indonesia (Moluccas)
29 C. theoblicki Yu & Li, 2019 ♂♀ China (Yunnan)
30 C. tongi Yu & Li, 2019 ♂♀ China (Yunnan)
31 C. zhengi Yu & Li, 2019 ♂♀ China (Yunnan)
32 C. mii Yu & Li, sp. nov. China (Yunnan)
33 C. subtongi Yu & Li, sp. nov. China (Yunnan)

Key to C. ternatensis group species occurring in Xishuangbanna (males)

Male of C. mii sp. nov. is unknown.

1 Palp with two tibial apophyses (Figs 57E, 67E) C. zhengi
Palp with single tibial apophysis 2
2 Emblous short, tip extending to 1/3 tegulum (Figs 57C, 67C) C. subkuu
Emblous distinctly long, tip extending basad more than 4/5 × length of tegulum (Figs 57A, B, D, 67A, B, D) 3
3 Tegular hump nearly quadrate (Figs 57A) C. theoblicki
tegular hump with a blunt and semi-circular tip, resembling a wave crest in ventral view (Figs Fig. 57B, D) 4
4 Embolar apex terminating at ca. 5 o’clock position of tegulum; tegular hump is ca. 1/3 tegulum length; tegular base with a papilliform flange (Figs 57B, 67B) C. tongi
Embolar tip terminating at approximately 4 o’clock position; tegular hump ca. 1/5 tegulum length; tegular base is unmodified (Figs 57D, 67D) C. subtongi sp. nov.

Key to C. ternatensis group species occurring in Xishuangbanna (females)

C. heteroducta is excluded due to lack of specimens.

1 Epigyne with atrium (Figs 77E, 85E, 93E) C. zhengi
Epigyne without atrium 2
2 Copulatory ducts short, not longer than epigyne length, not convoluted (Fig. 93A, B) 3
Copulatory ducts long, > 2 × longer than epigyne length, strongly convoluted (Fig. 93C, D) 4
3 Epigynal ridges triangular (Figs 77A, 85A) C. mii sp. nov.
Epigynal ridges more or less blade-shaped (Figs 77B, 85B) C. subkuu
4 Epigynal ridges diagonal (Figs 77C, 85C); anterior part of copulatory ducts forming 2 longitudinal loops (Fig. 93C) C. theoblicki
Epigynal ridges longitudinal (Figs 77D, 85D); anterior part of copulatory ducts with 2 transversal loops (Fig. 93D) C. tongi

Clubiona heteroducta Zhang & Yin, 1998

Clubiona heteroducta Zhang & Yin, 1998: 12, figs 12, 13 (♂♀).

Material examined

None.

Diagnosis and description

See Zhang and Yin (1998).

Distribution

China (Yunnan).

Clubiona mii Yu & Li, sp. nov.

Figs 27, 77A, 85A, 93A

Holotype

♀ (IZCAS-Ar 34758, YHCLU0065), China, Yunnan Province: Xishuangbanna: Mengla County: Nanshahe Village: monsoon forest, 21°36.388'N, 101°34.247'E, ca. 797 m, 13.VII.2012, Q.Y. Zhao and C.X. Gao leg.

Etymology

This species is named after Mr. Xiaoqi Mi (Tongren City, China) who has helped us greatly with this research.

Diagnosis

The female of the new species is easily distinguished from those of the other species in the group, with the exception of C. hystrix (Deeleman-Reinhold 2001: 103, figs 17, 18), by the general shape of the vulva but can be recognised by the: (1) triangular epigynal ridge (Figs 27A–C, 77A, 85A) (vs. pocket-like; Deeleman-Reinhold 2001 fig. 17); (2) copulatory openings close together (Figs 27A–C, 77A, 85A) (vs. copulatory openings separated by one diameter; Deeleman-Reinhold 2001 fig. 17).

Description

Female. Holotype (Fig. 27F, G): total length 2.63; carapace 1.29 long, 0.99 wide; opisthosoma 1.34 long, 0.90 wide. Carapace light orange, darker anteriorly, without distinct pattern, pars cephalica slightly narrowed, cervical groove indistinguishable; tegument smooth, all setae detached in ethanol. Eyes: AER slightly recurved, PER slightly wider than AER, almost straight in dorsal view. Eye sizes and interdistances: AME 0.08, ALE 0.07, PME 0.07, PLE 0.06, AMEAME 0.04, AMEALE 0.02, PMEPME 0.17, PMEPLE 0.05, MOQL 0.18, MOQA 0.22, MOQP 0.31. Chelicerae coloured as ocular area, with six promarginal and two retromarginal teeth. Sternum yellowish white, 1.38 long, 0.95 wide. Labium and endites coloured as chelicerae. Legs coloured as sternum, without markings. Leg measurements: I 1.84 (0.53, 0.73, 0.33, 0.25), II 1.98 (0.58, 0.77, 0.40, 0.23), III 1.81 (0.58, 0.59, 0.41, 0.22), IV 2.85 (0.83, 1.00, 0.71, 0.30). Abdomen yellowish white, uniformly coloured, clothed with dense setae, without pattern.

Epigyne (Figs 27A–E, 77A, 85A, 93A). Epigynal plate slightly longer than wide, spermathecae and bursae prominently visible through integument in ventral view, posterior margin not rebordered. Copulatory openings small, contiguous, situated at medial portion of epigynal plate posterior margin, hidden in two transverse ridges in ventral view. Ridge represented by triangular sclerite. Copulatory ducts thick and straight, close together, ascending parallel, entering the connecting piece located inside of bursal surface, then continuing upward and turning sideways, finally connecting to anteriorly located spermathecae. Spermathecae globular, separated by one diameter. Fertilisation ducts small, acicular. Bursae oblong, large, approximately as long as copulatory ducts, separated by 0.5 × diameters, ca. 1.5 × longer than wide, with smooth surface.

Male. Unknown.

Comments

According to WSC (2021), a total of seven described C. ternatensis group species are known only from males (See Table 4). Among them, C. bachmaensis, C. esuriens, and C. kuu were found form the adjacent area of Xishuangbanna. We cannot rule out the possibility that these three species are conspecific to C. mii sp. nov.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona subkuu Yu & Li, 2019

Figs 57C, 67C, 77B, 85B, 93B

Clubiona subkuu Yu & Li, 2019a: 164, figs 9A–E, 10A–H (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34604), 1♀ (paratype, IZCAS Ar 34605), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Lvshilin Forest Park, limestone tropical seasonal rainforest, 21°54.555'N, 101°16.860'E, ca. 610 m, 29.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♂ (YHCLU0039), XTBG, G213 roadside, leprosy village, 21°53.593'N, 101°17.329'E, ca. 559 m, 5.VIII.2018, H. Yu et al. leg; 1♀ (YHCLU0038), XTBG, secondary tropical forest, 21°54.833'N, 101°16.781'E, ca. 575 m, 31.VII.2018, Z.G. Chen leg.

Diagnosis and description

See Yu and Li (2019a). Male palp as in Figs 57C, 67C, epigyne as in Figs 77B, 85B, 93B.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona kuu.

Clubiona subtongi Yu & Li, sp. nov.

Figs 28, 29, 57D, 67D

Holotype

♂ (IZCAS-Ar 34759, YHCLU0056), : Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Anogeissus acuminata plantation, 21°54.033'N, 101°16.900'E, ca. 606 m, 2.VIII.2018, Z.G. Chen et al. leg.

Etymology

The specific name is taken from its similarity to Clubiona tongi; modified noun (name) in genitive case..

Diagnosis

Clubiona subtongi sp. nov. resembles C. tongi (Yu and Li 2019b: 212, figs 9A–E, 10E, F; Figs 57B, 67B) by having similar pale colouration and habitus (Fig. 29; Yu and Li 2019b: fig. 10E, F) and general shape of the palp (Figs 28A–E, 57B, 67B; Yu and Li 2019b: 212, fig. 9A–E) but can be distinguished by the: (1) embolar apex terminating at approximately the 4 o’clock position (Figs 28D, 57D) (vs. relatively longer tip terminating at approximately the 5 o’clock position; Figs 57B, 67B); (2) tegular hump that is ca. 1/5 tegulum length (Figs 28D, 57D) (vs. tegular hump ca. 1/3 tegulum length; Fig. 57B); (3) tegular base is unmodified (Figs 28B, 57D, 67D) (vs. with a papilliform flange; Fig. 67B).

Description

Male. Holotype (Fig. 29): Total length 4.63; carapace 2.13 long, 1.54 wide; opisthosoma 2.50 long, 1.16 wide. Carapace light brown, slightly darker on the front ridge, without pattern, ocular area slightly narrowed, cervical groove indistinct; tegument smooth, marginally clothed with long, thin setae. Eyes: in dorsal view, AER slightly recurved, PER slightly procurved, PER slightly wider than AER. Eye sizes and interdistances: AME 0.09, ALE 0.10, PME 0.12, PLE 0.11, AMEAME 0.06, AMEALE 0.03, PMEPME 0.26, PMEPLE 0.15, MOQL 0.33, MOQA 0.30, MOQP 0.46. Chelicerae brownish red, promargin with six teeth, retromargin with three teeth. Sternum yellowish white, 1.38 long, 0.95 wide. Labium and endites coloured as carapace. Legs yellowish white, without distinct markings. Leg measurements: I 4.28 (1.30, 1.70, 0.80, 0.48), II 4.70 (1.29, 1.98, 0.93, 0.50), III 3.83 (1.13, 1.28, 0.94, 0.49), IV 6.22 (1.88, 2.05, 1.79, 0.49). Abdomen lanceolate, dorsally grey with a lengthwise white heart shaped mark, reaching posterior half; with a pair of muscle depressions located on both sides of heart-shaped mark; venter centrally with an inverted trapezoidal orange patch.

Palp (Figs 28, 57D, 67D). Femur and patella unmodified. Tibia short, ca. 1/3 of cymbium length, with single retrolateral apophysis; RTA small, ca. 1/3 palpal tibia length, with a thumb-like base and spine-like tip. Tegulum elongate, oval, relatively flat, 2.1 × longer than wide, sperm duct distinct, V-shaped; tegular hump prominent, ca. 1/5 of tegulum length. Embolus filiform, originating on the retrolateral flank (ca. 10–11 o’clock on tegulum), aligning clockwise along the tegular hump, apex filiform, terminating at ca. 4 o’clock position.

Female. Unknown.

Comments

According to the WSC (2021), a total of eight described C. ternatensis group species are known only from females (See Table 4). We describe this species based on the male, although it may be synonymised in future.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona theoblicki Yu & Li, 2019

Figs 57A, 67A, 77C, 85C, 93C

Clubiona quadrata Yu & Li, 2019a: 161, figs 7A–E, 8A–H (♂♀)

Clubiona theoblicki Yu & Li, 2019c: 40 (replacement name).

Material examined

Type. Holotype ♂ (IZCAS Ar 34568), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary tropical forest, 21°54.380'N, 101°16.815'E, ca. 627 m, 23.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♀, XTBG, bamboo plantation, 21°53.901'N, 101°16.884'E, ca. 568 m, 12.V.2019, Z.G. Chen et al. leg; 1♂ (YHCLU0092) and 1♀ (YHCLU0093), XTBG, 48th km landmark in Menglun Nature Reserve, 21°53.997'N, 101°16.957'E, ca. 593 m, 11.VIII.2011, G. Zheng et al. leg.

Diagnosis and description

See Yu and Li (2019a). Male palp as in Figs 57A, 67A, epigyne as in Figs 77C, 85C, 93C.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona tongi.

Clubiona tongi Yu & Li, 2019

Figs 57B, 67B, 77D, 85D, 93D

Clubiona tongi Yu & Li, 2019b: 212, figs 9A–E, 10A–H (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34705), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, garbage dump, secondary tropical forest, 21°54.380'N, 101°16.815'E, ca. 627 m, 23.XI.2009, G. Tang and Z.Y. Yao leg; 1♀ (paratype, IZCAS Ar 34707), XTBG, G213 roadside, bamboo plantation, 21°54.622'N, 101°16.955'E, ca. 581 m, 26.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♂ (YHCLU0055), XTBG, rubber-tea plantation, 21°55.239'N, 101°15.854'E, ca. 572 m, 28.VII.2018, Z.G. Chen et al. leg; 1♀ (YHCLU0095), XTBG, 48th km landmark in Menglun Nature Reserve, 21°53.997'N, 101°16.957'E, ca. 593 m, 11.VIII.2011, G. Zheng et al. leg.

Diagnosis and description

See Yu and Li (2019b). Male palp as in Figs 57B, 67B, epigyne as in Figs 77D, 85D, 93D.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona theoblicki.

Clubiona zhengi Yu & Li, 2019

Figs 57E, 67E, 77E, 85E, 93E

Clubiona zhengi Yu & Li, 2019a: 167, figs 11A–E, 12A–H (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34583), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Anogeissus acuminata plantation, 21°54.017N, 101°16.900E, ca. 561 m, 27.IV.2019, Z.G. Chen et al. leg; 1♀ (paratype, IZCAS Ar 34590), XTBG, Lvshilin Forest Park, 21°54.609'N, 101°17.090'E, ca. 643 m, 17.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♂ (YHCLU0042) and 1♀ (YHCLU0043), XTBG, Anogeissus acuminata plantation, 21°54.033N, 101°16.900E, ca. 606 m, 2.VIII.2018, Z.G. Chen et al. leg.

Diagnosis and description

See Yu and Li (2019a). Male palp as in Figs 57E, 67E, epigyne as in Figs 77E, 85E, 93E.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona jaegeri.

Clubiona japonicola group

Diagnosis and description

See Mikhailov (1995).

Composition and distribution

According to Mikhailov (1995), the group includes only two species: C. japonicola Bösenberg & Strand, 1906 widespread from the Far East of Russia to the Philippines and Indonesia, and C. yasudai Ono, 1991, endemic to Hokkaido, Japan.

Comments

Morphological characters and our unpublished molecular data (pers. obs.) strongly suggest a close relationship between C. japonicola and C. riparia L. Koch, 1866 (assigned to the lutescens group by Mikhailov (1995)).

Clubiona japonicola Bösenberg & Strand, 1906

Clubiona japonicola Bösenberg & Strand, 1906: 281, pl. 16, fig. 498 (♂♀); Ono and Hayashi 2009: 535, figs 50–52 (♂♀); Yin et al. 2012: 1101, figs 580a–e, 3-14a–b (♂♀); Wang et al. 2018: 325, fig. 10A–F (♂♀). For full list of taxonomic references see WSC (2021)

Material examined

None.

Diagnosis and description

See Huang and Chen (2012).

Distribution

China (from Jili and south to Yunnan); Russia (Far East); South Korea; Japan; Philippines; Indonesia.

Clubiona filicata group

Tolophus Thorell, 1891: 26 (type species T. submaculatus Thorell, 1891).

Japoniona Mikhailov, 1990: 443 (described as subgenus).

Clubiona: Deeleman-Reinhold 2001: 90 (synonymised Tolophus and Japoniona)

Clubiona japonica group: Deeleman-Reinhold 2001: 90

Diagnosis

See Mikhailov (1990) and Yu et al. (2017a).

Description

See Mikhailov (1995) and Deeleman-Reinhold (2001).

Composition and distribution

A list of the species of the japonica group was provided by Yu et al. (2017a). Combined with new taxonomic data in the present paper, we provide an updated list (Table 5).

Comments

Japoniona Mikhailov,1990 was described to accommodate the japonica group. Later, the subgenus Japoniona was synonymised by Deeleman-Reinhold (2001) with Clubiona. A long-forgotten genus, Tolophus Thorell, 1891: 26 (type species T. submaculatus Thorell, 1891, belonging to the japonica group), is currently considered a junior synonym of Clubiona. The monophyly of the group is supported by morphological characters and molecular data (pers. obs.). However, the exact placement of the group within Clubiona or Clubionidae remains to be investigated using integrative taxonomy. Tolophus should perhaps be removed from synonymy with Clubiona.

Table 5.

Clubiona filicata group.

Species name Known sex Distribution
1 C. annuligera Lessert, 1929 ♂♀ Congo, Mozambique
2 C. abnormis Dankittipakul, 2008 ♂♀ Thailand (Nakhorn Ratchasima), Laos, China (Yunnan)
3 C. bilobata Dhali, Roy, Saha & Raychaudhuri, 2016 India (West Bengal)
4 C. calycina Wu & Zhang, 2014 ♂♀ China (Henan)
5 C. campylacantha Dankittipakul, 2008 ♂♀ Thailand (Loei)
6 C. charleneae Barrion & Litsinger, 1995 ♂♀ Philippines (Quezon)
7 C. circulata Zhang et Yin, 1998 ♂♀ China (Yunnan)
8 C. coreana Paik, 1990 ♂♀ Russia (south part of the Far East), Korea China (Niaoning)
9 C. reichlini Schenkel, 1944 ♂♀ China (Zhejiang)
10 C. digitata Dankittipakul, 2012 ♂♀ Thailand (Loei, Pathum Thani)
11 C. drassodes O. P.-Cambridge, 1874 ♂♀ India, Bangladesh, China
12 C. filicata O. Pickard-Cambridge, 1874 ♂♀ Pakistan to Taiwan, south China, south to Thailand
13 C. filifera Dankittipakul, 2008 ♂♀ Thailand (Nakhorn Ratchasima)
14 C. filoramula Zhang & Yin, 1998 China (Yunnan)
15 C. gallagheri Barrion & Litsinger, 1995 Indonesia (Java)
16 C. grucollaris Yu, Zhang & Chen, 2017 ♂♀ China (Hainan, Guizhou, and Yunnan)
17 C. japonica L. Koch, 1878 ♂♀ Russia (Sakhalin, Kurile Is.), China (Taiwan), Korea, Japan
18 C. lala Jäger & Dankittipakul, 2010 Laos (China (Yunnan).
19 C. melanosticta Thorell, 1890 ♂♀ Thailand (Chiang Mai, Samut Songkram), Indonesia (Sumatra, Krakatau), New Guinea, Laos, China (Yunnan).
20 C. munda Thorell, 1887 Myanmar (Kachin State)
21 C. nigromaculosa Blackwall, 1877 ♂♀ Seychelles, Réunion
22 C. octoginta Dankittipakul, 2008 ♂♀ Thailand (Ubon Ratchathani)
23 C. picturata Deeleman-Reinhold, 2001 ♂♀ Indonesia (Bali)
24 C. pila Dhali, Roy, Saha & Raychaudhuri, 2016 India (West Bengal)
25 C. pupula Thorell, 1897 ♂♀ Myanmar (Kachin State)
26 C. scandens Deeleman-Reinhold, 2001 ♂♀ Malaysia (Borneo)
27 C. submaculata (Thorell, 1891) ♂♀ India (Nicobar Is.)
28 C. suthepica Dankittipakul, 2008 ♂♀ Thailand (Chiang Mai), China (Yunnan)
29 C. vigil Karsch, 1879 ♂♀ Kuril Isles, Korea, Japan, China (Hebei, Hubei)
30 C. yueya Yu & Li, 2019 ♂♀ China (Yunnan)
31 C. zhanggureni Yu & Li, 2019 China (Yunnan)
32 C. zhangyongjingi Li & Blick, 2019 China (Yunnan)
33 C. banna Yu & Li, sp. nov. ♂♀ China (Yunnan)

Key to C. filicata group species occurring in Xishuangbanna (males)

C. filoramula is excluded due to lack of specimens.

1 Conductor absent (Figs 58A, B, 68A, B) 2
Conductor large and beak-shaped, transversely aligned anteriorly (Figs 58C, 59A–D, 60A–D, 68C, 69A–D, 70A–D) 3
2 Embolus short, no longer than tegulum width, sickle-shaped; tegular apophysis absent (Figs 58A, 68A) C. reichlini
Conductor distinctly longer than tegulum width, filiform and slender, tapering gradually in ca. two loops; tegular apophysis small and digitiform (Figs 58B, 68B) C. filicata
3 Tegular apophysis distinct, heavily sclerotised (Figs 58C, 60A–D, 68C, 70A–D) 4
Tegular apophysis absent, or present but reduced, partly membranous, indistinct (Figs 59A–D, 69A–D) 8
4 Tegulum obscuring sperm duct in ventral view (Fig. 60C, D) 5
Sperm duct sinuate and distinct (Figs 58C, 60A, B) 6
5 Tegular apophysis large, crescent-shaped; embolus spiniform (Fig. 60D) C. yueya
Tegular apophysis small, beak-shaped; embolus filiform (Fig. 60C) C. abnormis
6 Embolar apex coiled (Fig. 42D); tegular apophysis large and boomerang shaped in ventral view (Fig. 60B) C. lala
Embolar tip not coiled, tegular apophysis not as above 7
7 Tegular apophysis with tubercle-shaped base and rostrate tip; sperm duct sinuate, forming a loop along tegular margin (Fig. 58C) C. banna sp. nov.
Tegular apophysis petal-shaped, sperm duct U- or S-shaped (Fig. 60A) C. grucollaris
8 Retrolateral tibial apophysis hook-shaped, strongly excavated on the ventral side; cymbium with basolateral extension (Fig. 69D) C. suthepica
Retrolateral tibial apophysis not so; cymbium without basolateral extension 9
9 Conductor strongly sclerotised, horn-shaped, pointing retrolatero-distally (Fig. 59C); retrolateral tibial apophysis bent, thumb-like (Fig. 69C) C. circulata
Conductor membranous except for the beak-shaped tip (Fig. 59A, B), retrolateral tibial apophysis triangular (Fig. 69A, B) 10
10 Embolus spiniform (Fig. 59B) C. zhanggureni
Embolus filiform (Fig. 59A) C. melanosticta

Key to females of C. filicata group species occurring in Xishuangbanna (females)

(the females of C. abnormis, C. filicata, C. filoramula, C. zhanggureni, and C. zhangyongjingi are excluded due to lack of specimens)

1 Atrium small, width no more than 1/2 epigyne width (Figs 78E, 86E) C. banna sp. nov.
Atrium broad, width almost equal to epigyne width (Figs 78F, 79A–F, 86F, 87A–F) 2
2 Posterior margin of atrium delimited 3
Posterior margin of atrium not delimited 4
3 Atrium anterior margin not delimited (Figs 78F, 86F); spermathecae consisting of base, stalk and head (Fig. 94F) C. melanosticta
Atrium anterior margin distinct and long, ˄-shaped (Figs 79A, 87A); spermathecae distinctly smaller than bursae, lobe-shaped, not subdivided (Fig. 95A) C. circulata
4 Atrium anterior margin heavily sclerotised, M-shaped, lateral atrial margins not rebordered (Figs 79E, 87E) C. suthepica
Atrium anterior margin not sclerotised; lateral atrial margins rebordered 5
5 Atrium anterior margin medially concave (Figs 79C, 87C, F) 6
Atrium anterior margin medially not concave (Figs 79B, 87B, D) 7
6 Spermathecae larger than bursae, consisting of papilliform base, tubular stalk and ovoid head, ascending spirally (Fig. 95C) C. grucollaris
Spermathecae distinctly smaller than bursae, consisting of bean-shaped proximal part and digitiform distal part (Fig. 95F) C. yueya
7 Atrium more rectangular (Fig. 87D); spermathecae located in the centre of vulva, well separated from atrium anterior margin; spermathecae separated by ca. one diameter, consisting of base and head (Fig. 95D) C. lala
Atrium more cambered (Fig. 79B, 87B); spermathecae located at anterior part of vulva, close to atrium anterior margin; spermathecae close together, consisting of base, stalk and head (Fig. 95B) C. reichlini

Clubiona abnormis Dankittipakul, 2008

Figs 30, 31, 60C, 70C

Clubiona abnormis Dankittipakul, in Dankittipakul and Singtripop 2008a: 44, figs 28, 29, 61–68 (♂♀); Jäger and Dankittipakul 2010: 32, figs 27, 31–33 (♀).

Material examined

1♂ (YHCLU0113), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, 48th km landmark in Menglun Nature Reserve, 21°58.704'N, 101°19.748'E, ca. 1088 m, 12.VIII.2011, G. Zheng et al. leg.

Diagnosis and description

See Dankittipakul and Singtripop (2008a). Male palp as in Figs 30, 60C, 70C, habitus as in Fig. 31.

Distribution

Thailand (Nakhorn Ratchasima), Laos, China (Yunnan).

Most similar species

Clubiona vigil.

Clubiona banna Yu & Li, sp. nov.

Figs 32, 33, 58C, 68C, 78E, 86E, 94E

Holotype

♂ (IZCAS-Ar 34760), China: Yunnan Province: Xishuangbanna: Mengla County: Xiaolongha Village, 21°24.798'N, 101°37.880'E, ca. 693 m, 28.VI.2012, Q.Y. Zhao and C.X. Gao leg; paratype: 1♀ (IZCAS-Ar 34761, YHCLU0139), Menglun Town, XTBG, primary tropical seasonal forest, 21°57.445'N, 101°12.997'E, ca. 774 m, 30.VIII.2007, G. Zheng leg. Other material examined. 1♂ (YHCLU0104), same data as holotype.

Etymology

The species name is derived from the name of the type locality; noun in apposition.

Diagnosis

Males of C. banna sp. nov. can be easily distinguished from those of all others in the species group by the tegular apophysis, having a tubercle-shaped base and a rostrate tip (Fig. 32B, D, E). Females of C. banna sp. nov. are similar to those of C. vigil (Kim and Lee 2014: 38, fig. 26B) by the relatively small atrium (vs. atria usually broad in almost all members of the C. filicata group, including C. circulata, C. reichlini, C. grucollaris and C. lala; Figs 35A, B, 79A, 87A, 37A, B, 79B, 87B, 41A, B, 79C, 87C, 43A, B, 79D, 87D) but can be recognised by the atrium being anteriorly elliptic and posteriorly shaped like an inverted triangle (Figs 33A, B, 78E, 86E) (vs. cordiform in C. vigil).

Description

Male. Holotype (Fig. 33E, F): Total length 6.53; carapace 2.84 long, 2.02 wide; opisthosoma 3.69 long, 1.73 wide. Carapace brown, marginally dark, a pair of Y-shaped black markings starting from behind PME and PLE, almost reaching reddish fovea; pars cephalica distinctly narrowed; cervical groove and radial grooves indistinct; tegument smooth, marginally clothed with short, dense setae. Eyes: in dorsal view, both anterior and posterior eye rows recurved, PER slightly wider than AER. Eye sizes and interdistances: AME 0.11, ALE 0.15, PME 0.15, PLE 0.14, AMEAME 0.14, AMEALE 0.07, PMEPME 0.26, PMEPLE 0.17, MOQL 0.40, MOQA 0.36, MOQP 0.53. Chelicerae brown, with red fangs, with three promarginal and two retromarginal teeth. Sternum centrally pale brown, marginally dark, 1.52 long, 1.02 wide. Labium and endites coloured as chelicerae. Legs light brown, without distinct markings. Leg measurements: I 9.08 (2.61, 3.60, 1.85, 1.02), II 10.55 (2.87, 3.96, 2.59, 1.13), III 8.25 (2.44, 2.65, 2.46, 0.71), IV 10.98 (3.02, 3.50, 3.43, 1.02). Abdomen: dorsum with broken dark median band in anterior half, posteriorly with 5 chevrons; venter with three dark longitudinal lines.

Palp (Figs 32A–E, 58C, 68C). Tibia short, ca. 1/3 of cymbium length, with single retrolateral apophysis; hammer-like or clavate RTA small, slightly curved, and bluntly pointed. Tegulum more or less spherical, 2.1 × longer than wide, proapically and apically membranous, slightly excavated prolatero-apically to accommodate embolus; sperm duct sinuate, forming a loop along tegular margin. Embolus filiform, arising at approximately the 9–10 o’clock position, terminating at ca. 12 o’clock position, tip hidden behind conductor. Conductor large, beak-shaped, transversely aligned at the apical portion of the bulb, basal part partly membranous, terminal part heavily sclerotised, directed retrolaterad then abruptly bending distad. Tegular apophysis with tubercle-shaped base and rostrate tip, located at distal-retrolateral position of tegulum (ca. 1 o’clock position of tegulum).

Female. Paratype (Fig. 33G, H). Total length 8.05; carapace 3.40 long, 2.19 wide; opisthosoma 4.65 long, 2.95 wide. Eye sizes and interdistances: AME 0.15, ALE 0.20, PME 0.18, PLE 0.16, AMEAME 0.11, AMEALE 0.11, PMEPME 0.31, PMEPLE 0.20, MOQL 0.49, MOQA 0.45, MOQP 0.66. Chelicerae with three promarginal and two retromarginal teeth. Sternum 1.88 long, 1.14 wide. Leg measurements: I 8.71 (2.42, 3.50, 1.78, 1.01), II 9.56 (2.67, 3.71, 2.10, 1.07), III 7.78 (2.23, 2.75, 2.08, 0.72), IV 10.60 (2.86, 3.47, 3.21, 0.97). Colouration lighter than in male. Other characters as in male.

Epigyne (Figs 33A–D, 78E, 86E, 94E). Epigynal plate nearly square, margin not rebordered, spermathecae and bursae indistinctly visible through integument. Atrium small, with delimited margin, ca. 1/2 epigyne length and 1/3 epigyne width, anteriorly elliptic, posteriorly triangular. Copulatory openings located at lateral atrial borders. Copulatory ducts short, running sideways, then retracing posteriorly to bursae. Spermathecae with subglobular proximal part and tubular distal part; the two proximal parts separated by 0.5 diameters, and the two distal parts close together. Fertilisation ducts short and curved, acicular, located on distal surface of spermathecae. Bursae reniform, ca. 1.5 × longer than wide, close together, surface membranous and smooth.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona circulata Zhang & Yin, 1998

Figs 34, 35, 59C, 69C, 79A, 87A, 95A

Clubiona circulata Zhang & Yin, 1998: 9, figs 1–2 (♀ only, ♂ mismatched).

Clubiona vukomi Jäger & Dankittipakul, 2010: 27, figs 13–21 (♂). Syn. nov.

Material examined

1♂, China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, rubber plantation, 21°54.498'N, 101°16.326'E, ca. 586 m, 17.VII.2007, G. Zheng leg; 2♀, XTBG, rubber plantation, 21°54.350'N, 101°16.461'E, ca. 614 m, 11.VIII.2007, G. Zheng leg; 1♂ (YHCLU0108), Jinghong City: Menghai County: Manda Village, secondary forest, 22°1.702'N, 100°23.697'E, ca. 1188 m, 28.VII.2012, Q.Y. Zhao and Z.G. Chen leg; 1♀ (YHCLU0156), XTBG, seedling culture base, 21°54.007'N, 101°16.395'E, ca. 550 m, 10.V.2019, H. Yu et al. leg; 7♂ 6♀, XTBG, Flocculus banyan plantation, 22°4.598'N, 100°37.013'E, ca. 1137 m, 21.VIII.2011, Q.Y. Zhao and C.X. Gao leg;

Diagnosis

The female of C. circulata is easily differentiated from other members of the group by having an epigynal atrium with ˄-shaped anterior margin and a V-shaped posterior margin (Figs 35A, B, 79A, 87A) (vs. anterior margin not ˄-shaped, atrial posterior margin absent in almost all others, including C. reichlini, C. grucollaris, and C. lala; Figs 37A, B, 79B, 87B, 41A, B, 79C, 87C, 43A, B, 79D, 87D). Males of C. cicrulata differ from all other group members by the large and strongly sclerotised, horn-shaped conductor, pointing retrolatero-distally (Fig. 34B–E) (vs. conductors of almost all other congroupers, such as C. grucollaris and C. suthepica, are beak-shaped, pointing retrolatero-proximally; Figs 40B–E, 60A, 70A, 46B–E, 59D, 69D).

Description

Male. See Jäger and Dankittipakul (2010). Male palp as in Figs 34, 59C, 69C, habitus as in Fig. 35E, F.

Female. (Fig. 35G, H): Total length 6.00; carapace 2.67 long, 2.01 wide; opisthosoma 3.33 long, 1.59 wide. Carapace reddish brown, anteriorly darker, with a pair of indistinct, Y-shaped purplish patterns starting from behind PER, almost reaching indistinct cervical groove, fovea dark reddish; cephalic region raised, radial grooves indistinct; tegument smooth, with erect, thin, dark setae on the front ridge. Eyes: AER slightly recurved, PER slightly wider than AER, almost straight in dorsal view. Eye sizes and interdistances: AME 0.10, ALE 0.14, PME 0.13, PLE 0.09, AMEAME 0.08, AMEALE 0.04, PMEPME 0.24, PMEPLE 0.16, MOQL 0.40, MOQA 0.36, MOQP 0.51. Chelicerae robust and dark brownish red, promargin with three teeth, retromargin with two subequal teeth. Sternum pale yellow, 1.43 long, 1.01 wide. Labium and endites orange. Legs light brown, without distinct markings. Leg measurements: I 7.40 (2.11, 3.00, 1.46, 0.83), II 7.88 (2.26, 3.14, 1.62, 0.86), III 6.30 (1.69, 2.23, 1.88, 0.53), IV 9.58 (2.55, 3.34, 2.96, 0.74). Abdomen elongate, oval, with a thick tuft of setae anteriorly; dorsum with broken purplish longitudinal band medially, starting anteriorly for half the length; 6–7 pair of broken lateral bands fused posteriorly. Venter uniformly creamy white, without markings.

Epigyne (Figs 35A–D, 79A, 87A, 95A). Epigynal plate ca. 1.1 × longer than wide, through which large spermathecae and bursae are clearly visible. Atrium distinctly large, with rebordered margin, more than 2/3 epigyne length and 4/5 epigyne width; atrial anterior margin long and shaped like ˄, atrial posterior margin relatively short and nearly V- or U-shaped. Copulatory openings near the middle part of the epigyne, close to the anterolateral borders of the APM. Copulatory ducts distinct, almost equal to bursae length, extending obliquely toward the anterior, between the spermathecae, before abruptly bending posteriorly, finally entering the connecting piece between the spermathecae and bursae. Spermathecae relatively small, lobe-shaped, separated by two diameters. Fertilisation ducts acicular, membranous, located terminally on spermathecae. Bursae oblong, 1.4 × longer than wide, close together, bursal surface hyaline, wrinkled and ribbed, inside pigmented and sclerotised.

Remarks

Clubiona circulata was described based on five females and two males from Xishuangbanna. The female was chosen as the holotype. However, we have found that the male and female of C. circulata were mismatched. While examining spider specimens collected from Xishuangbanna, we found pairs of filicata group specimens in the same location that have a similar habitus, markings, leg spination, and other characters (Fig. 35E–H). Therefore, it is very likely they are the opposite sexes of the same species. The females were identified as C. circulata based on comparison with the original illustrations of Zhang and Yin (1998). However, the males were identified to be C. vukomi, which was established as a new species by Jäger and Dankittipakul (2010) from Luang Nam Tha Province in Laos. Our molecular analysis of COI indicates that the female of C. circulata and the male of C. vukomi are conspecific, and therefore C. vukomi should be considered a junior synonym of C. circulata.

Distribution

Thailand (Chiang Mai Province and district, Chai Ya Phum Province), Laos (Luang Nam Tha Province), China (Yunnan).

Clubiona reichlini Schenkel, 1944, species resurrected

Figs 36, 37, 58A, 68A, 79B, 87B, 95B

Clubiona reichlini Schenkel, 1944: 203, fig. 14 (♂♀).

Material examined

1♂ (YHCLU0263), 1♀(YHCLU0264), China: Yunnan Province: Xishuangbanna: Mengla County: Xiaolongha Village, 22°5.017'N, 100°22.084'E, ca. 1118 m, 24.VII.2012, Q.Y. Zhao and Z.G. Chen leg.

Diagnosis and description

See Schenkel (1944). Male palp as in Figs 36, 58A, 68A, epigyne as in Figs 37A–D, 79B, 87B, 95B, habitus as in Fig. 37E–H.

Remarks

Clubiona reichlini was considered a senior synonym of C. deletrix O. Pickard-Cambridge, 1885 by Zhang (1991), but this is not accepted here based on comments by Y. Marusik (pers. comm.). As a result, C. reichlini is removed from synonymy with C. deletrix.

Distribution

China (Zhejiang, Yunan).

Most similar species

Clubiona campylacantha.

Clubiona filicata O. Pickard-Cambridge, 1874

Figs 38, 39, 58B, 68B

Clubiona filicata O. Pickard-Cambridge, 1874: 413, pl. 52, fig. 35 (♂♀); Dankittipakul et al. 2012: 59, figs 25–31 (♂♀); Caleb 2020: 15719, figs 4A–F, 25G (♀).

Clubiona distincta Thorell, 1887: 48 (♀).

Clubiona swatowensis Strand, 1907: 562 (♀); Strand 1909: 39, fig. 24 (♀).

Clubiona pashabhaii Patel & Patel, 1973: 2, fig. 1a–c (♀).

Clubiona foliata Keswani & Vankhede, 2014: 36, figs 1–13 (♂♀). For full list of taxonomic references see WSC (2021).

Material examined

1♂ (YHCLU0107), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, flower garden, 21°55.919'N, 101°14.994'E, ca. 545 m, 14.V.2019, C. Wang and H. Yu leg.

Diagnosis and description

Male. See Dankittipakul and Singtripop (2008a). Male palp as in Figs 38, 58B, 68B, habitus as in Fig. 39.

Female. See Keswani and Vankhede (2014).

Remarks

Clubiona deletrix was described based on both sexes in the original publication. Y. Marusik studied the types of C. deletrix and C. filicata and the original drawings. He found that the male and female of C. deletrix were not conspecific in the original description, and the male type is C. filicata (pers. comm.).

Distribution

From Pakistan to Taiwan, south to Thailand, China (Fujian, Hunan, Guangdong, Guangxi, Taiwan, Yunnan).

Most similar species

Clubiona filoramula.

Clubiona filoramula Zhang & Yin, 1998

Clubiona filoramula Zhang & Yin, 1998: 12, figs 9–11 (♂).

Material examined

None.

Diagnosis and description

See Zhang and Yin (1998).

Distribution

China (Yunnan).

Most similar species

Clubiona filicata.

Clubiona grucollaris Yu, Zhang & Chen, 2017

Figs 40, 41, 60A, 70A, 79C, 87C, 95C

Clubiona grucollaris Yu et al., 2017a: 4, figs 1–2, 4–6, 13–19 (♂♀).

Material examined

1♂ (YHCLU0105), 1♀ (YHCLU0106), China: Yunnan Province: Xishuangbanna: Jinghong City: Menga Town: Wengnan Village: secondary forest, 22°4.997'N, 100°22.223'E, ca. 1137 m, 25.VII.2012, Q.Y. Zhao and Z.G. Chen leg.

Diagnosis and description

See Yu et al. (2017a). Male palp as in Figs 40, 60A, 70A, epigyne as in Figs 41A–D, 79C, 87C, 95C, habitus as in Fig. 41E–H.

Distribution

China (Hainan, Guizhou, and Yunnan).

Most similar species

Clubiona lala.

Clubiona lala Jäger & Dankittipakul, 2010

Figs 42, 43, 60B, 70B, 79D, 87D, 95D

Clubiona lala Jäger & Dankittipakul, 2010: 29, figs 22–25, 28–30 (♀).

Material examined

1♂, China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, tropical evergreen rainforest, 21°55.139'N, 101°16.295'E, ca. 523 m, 30.XI.2009, G. Tang and Z.Y. Yao leg; 1♀, XTBG, rubber plantation, 21°54.554N, 101°16.311'E, ca. 570 m, 14.V.2019, Z.G. Chen leg; 1♂ (YHCLU0110), Mengla County: Xiaolongha Village, 21°24.159'N, 101°37.178'E, ca. 635 m, 14.V.2019, Q.Y. Zhao and C.X. Gao leg; 1♀ (YHCLU0111), Jinghong City: Mengla County: Bubang Village, 21°36.384'N, 101°34.543'E, ca. 823 m, 10.VII.2012, Q.Y. Zhao and C.X. Gao leg.

Diagnosis

The male of C. lala resembles those of C. grucollaris (Figs 40, 60A, 70A) in having a long, columnar conductor base and beak-shaped conductor apex but differs in the following: the embolar apex is coiled (Figs 42D, E, 60B) (vs. not coiled; Figs 40, 60A); the tegular apophysis is boomerang-shaped in ventral view (Figs 42D, 60B) (vs. petal-shaped; Figs 40B, D, E, 60A, 70A); the finger-like retrolateral tibial apophysis (Figs 42A, B, 70B) (vs. triangular; Figs 40A, B, 70A). The female of C. lala can be separated from that of C. grucollaris by the atrium anterior margin medially not concave (vs. medially concave) (cf. Fig. 87C and 87D), spermathecae tubular, consisting of base and head (vs. ascending spirally, consisting of base, stalk, and head) (cf. Fig. 95C and 95D). The female of C. lala also appears to be closely related to C. campylacantha (Dankittipakul and Singtripop 2008a: 38, figs 2–4, 13, 14, 38–40), C. octoginta (Dankittipakul and Singtripop 2008a: 39, figs 18–19, 45–47) and C. reichlini (Figs 37A–D, 79B, 87B) by the general shape of the atrium and vulva but can be easily distinguished from these species by the: (1) more rectangular atrium (Figs 43A, B, 87D) (vs. atrium more cambered); (2) spermathecae posterior to the atrium, well separated from atrium anterior margin (Figs 43C, D, 95D) (vs. spermathecae situated anteriorly, close to atrium anterior margin); (3) spermathecae separated by ca. one diameter (Figs 43C, D, 95D). (vs. spermathecae close together).

Description

Male. (Fig. 43E, F): Total length 6.30; carapace 2.95 long, 1.96 wide; opisthosoma 3.53 long, 1.53 wide. Carapace brown, distinctly dark brown in ocular area, with a distinct pattern on pars cephalica consisting of a pair of dark, lateral bands and Ψ-shaped markings behind PER; ocular region distinctly narrowed, cervical groove and radial grooves indistinguishable; tegument smooth, clothed with short, dense setae. Eyes: AER slightly recurved, PER slightly procurved, the former wider than the latter. Eye sizes and interdistances: AME 0.11, ALE 0.15, PME 0.12, PLE 0.13, AMEAME 0.12, AMEALE 0.10, PMEPME 0.27, PMEPLE 0.16, MOQL 0.36, MOQA 0.35, MOQP 0.52. Chelicerae robust and dark brown, dorsally with dark pattern. Cheliceral furrow with three anterior and two posterior teeth. Sternum yellowish white, 1.47 long, 0.96 wide. Labium and endites light orange. Legs brownish, all legs with conspicuous dark brown annuli on the distal parts of the femur, patella, tibia, metatarsus, and tarsus. Leg measurements: I 6.51 (1.78, 2.67, 1.32, 0.75), II 6.99 (1.90, 2.80, 1.44, 0.86), III 5.62 (1.72, 1.60, 1.74, 0.56), IV 8.29 (2.44, 2.93, 2.29, 0.60). Abdomen brown, with conspicuous anterior setal tufts; dorsum light yellow, antero-laterally with disconnected longitudinal bands, posteriorly with dark purple markings; venter with two indistinct purplish longitudinal markings.

Palp (Figs 42A–E, 60B, 70B). Tibia short, ca. 1/2 × cymbium length, with retrolateral apophysis; RTA digitiform, broad at base, apex truncated. Bulb more or less spherical, ca. twice longer than wide, oval; sperm duct sinuate, running an irregular course in the postero-retrolateral part of the tegulum. Embolus flagelliform; embolar base situated meso-prolaterally on the tegulum; embolar apex coiled, resting on an apical portion of the tegulum, covered by conductor in prolateral view. Conductor large, longer than 1/2 length of tegulum, with a heavily sclerotised and beak-shaped apex, base membranous, long, and columnar. Tegular apophysis heavily sclerotised, boomerang-shaped in ventral view.

Female. See Jäger and Dankittipakul (2010). Epigyne as in Figs 43A–D, 79D, 87D, 95D, habitus as in Fig. 43G–H.

Distribution

Laos, China (Yunnan).

Remarks

Male of the species is described for the first time.

Clubiona melanosticta Thorell, 1890

Figs 44, 45, 59A, 69A, 78F, 86F, 94F

Clubiona melanosticta Thorell, 1890: 374 (♂); Thorell 1895: 42 (♀); Deeleman-Reinhold 2001: 123, figs 51–52 (♂); Dankittipakul and Singtripop 2008a: 42, figs 8–10, 52–54 (♂).

Clubiona melanothele Thorell, 1895: 42 (♀). Syn. nov.

Material examined

1♂3♀, China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Paramichelia baillonii plantation (ca. 20 yr.), 21°54.200'N, 101°16.923'E, ca. 608 m, 18.VIII.2007, G. Zheng leg; 1♂ (YHCLU0011), XTBG, teak plantation, 21°54.117'N, 101°16.167'E, ca. 549 m, 8.VIII.2018, H. Yu et al. leg; 1♀ (YHCLU0164), XTBG, 100 acre-feet sample plot (beside a hut), 21°54.117'N, 101°16.167'E, ca. 549 m, 11.VIII.2018, H. Yu et al.

Diagnosis and description

See Deeleman-Reinhold (2001). Male palp as in Figs 44, 59A, 69A, epigyne as in Figs 45A–D, 78F, 86F, 94F, habitus as in Fig. 45E–H.

Remarks

Clubiona melanosticta and C. melanothele were considered separate species for more than 120 years. After examining the holotypes, Deeleman-Reinhold (2001) illustrated the two species and suggested that they could be conspecific; however, she made no taxonomic changes at the time. New material has been collected from Xishuangbanna containing both sexes. According to drawings of Deeleman-Reinhold (2001), the males were identified as C. melanosticta, and the females were identified as C. melanothele. Based on morphology (Fig. 45E–H) and DNA barcoding data (Table 1), we matched the females and males together. Therefore, the two names are synonymised, and priority is given to C. melanosticta.

Distribution

Thailand (Chiang Mai, Samut Songkram), Indonesia (Sumatra, Krakatau), New Guinea, Myanmar, Laos, China (Yunnan).

Most similar species

Clubiona zhanggureni.

Clubiona suthepica Dankittipakul, 2008

Figs 46, 47, 59D, 69D, 79E, 87E, 95E

Clubiona suthepica Dankittipakul, in Dankittipakul and Singtripop 2008a 42, figs 22, 23, 55–58 (♂ only, ♀ mismatched).

Material examined

1♂, China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary tropical montane evergreen broad-leaved forest, 21°57.534'N, 101°12.300'E, ca. 860 m, 4.VIII.2007, Guo Zheng leg; 1♂ (YHCLU0114), XTBG, 48th km landmark in Menglun Nature Reserve, 21°58.704'N, 101°19.748'E, ca. 1088 m, 12.VIII.2011, G. Zheng et al. leg; 1♀ (YHCLU0209), XTBG, 48th km landmark in Menglun Nature Reserve, 21°58.764'N, 101°19.748'E, ca. 1038 m, 10.VIII.2011, Q.Y. Zhao and Z.G. Chen leg.

Diagnosis

Females of C. suthepica can be easily distinguished from other members of the group by the heavily sclerotised anterior margin of the atrium (Figs 47A, B, 79E, 87E). The male of C. suthepica differs from other members of the group by having a hook-shaped retrolateral tibial apophysis (Figs 46B, 69D) (vs. retrolateral tibial apophysis variable but not hook-shaped; for example, triangular in C. melanosticta and C. zhanggureni, hammer-like or clavate in C. banna sp. nov., digitiform in C. lala and C. yueya.; Figs 68C, 69A, B, 70B, D), the conductor apex terminating at ca. 9 o’clock position (Figs 46B, 59D) (vs. relatively shorter tip terminating at ca. 7–8 o’clock position in other species of the group; Figs 58C, 59A–C, 60A–D).

Description

Male. See Dankittipakul and Singtripop (2008a). Palp as in Figs 46, 59D, 69D, habitus as in Fig. 47E, F.

Female. (Fig. 47G, H): Total length 6.46; carapace 2.63 long, 1.88 wide; opisthosoma 3.83 long, 2.16 wide. Carapace brown, distinctly dark brown in ocular area, with a distinctive pattern on pars cephalica consisting of a pair of dark lateral bands and Ψ-shaped markings behind PER; ocular area slightly narrowed, cervical groove and radial grooves indistinguishable; tegument smooth, clothed with short setae. Eyes: AER slightly recurved, PER slightly wider than AER, almost straight in dorsal view. Eye sizes and interdistances: AME 0.14, ALE 0.15, PME 0.14, PLE 0.13, AMEAME 0.13, AMEALE 0.10, PMEPME 0.28, PMEPLE 0.20, MOQL 0.39, MOQA 0.37, MOQP 0.54. Chelicerae robust and dark brownish red, cheliceral furrow with three anterior and two posterior teeth. Sternum light yellow, 1.02 long, 0.68 wide. Labium and endites orange. Legs light yellow, femora with a broad distal band occupying almost half its length; tibiae with broad distal and proximal annuli; metatarsi with dark, thin distal annulus; tarsi pale yellow. Leg measurements: I 6.21 (1.83, 2.44, 1.17, 0.77), II 6.43 (1.89, 2.44, 1.28, 0.83), III 5.37 (1.65, 1.80, 1.34, 0.59), IV 7.35 (1.92, 2.57, 2.14, 0.73). Abdomen oval, with conspicuous anterior setal tufts, dorsum with dense grey setae and a broken purplish median band, half opisthosoma length, posteriorly with paired purplish markings consisting of numerous stripes and spots; venter yellowish white, medially with a longitudinal and linear marking.

Epigyne (Figs 47A–D, 79E, 87E, 95E). Epigynal plate nearly square, copulatory ducts visible through transparent integument, ca. 1/3 epigyne width. Anterior margin (or hood) heavily sclerotised, M-shaped, distinctly wide, almost equal to epigyne width. Copulatory openings indistinct, located in the hood. Copulatory ducts relatively long, nearly equal to bursal diameter. Spermathecae consisting of fan-shaped head and lobe-shaped base, with small fertilisation ducts terminally; the two spermathecal bases separated by 1.2 × length. Bursae close together, more or less spherical, surface translucent and wrinkled.

Distribution

Thailand (Chiang Mai), China (Yunnan).

Remarks

The female of the species is described for the first time.

Clubiona yueya Yu & Li, 2019

Figs 60D, 70D, 79F, 87F, 95F

Clubiona yueya Yu & Li, 2019b: 215, figs 11A–E, 12A–G (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34709), 1♀ (paratype, IZCAS Ar 34711), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, 300 acre-feet bamboo plantation, 21°53.901'N, 101°16.884'E, ca. 515 m, 7.VIII.2018, H. Yu and Z.G. Chen leg. Other material examined. 1♂ (YHCLU0116) and 1♀ (YHCLU0117), same locality as holotype, 12.V.2019, Z.G. Chen et al. leg.

Diagnosis and description

See Yu and Li (2019b). Male palp as in Figs 60D, 70D, epigyne as in Figs 79F, 87F, 95F.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona lala.

Clubiona zhanggureni Yu & Li, 2019

Figs 59B, 69B

Clubiona zhanggureni Yu & Li, 2019b: 216, figs 13A–E, 14A–C (♂).

Material examined

Types. Holotype ♂ (IZCAS Ar 34714), 1♂ (paratype, IZCAS Ar 34715), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary tropical montane evergreen broad-leaved forest, 21°57.809'N, 101°12.173'E, ca. 888 m, 4.VIII.2007, G. Zheng leg.

Diagnosis and description

See Yu and Li (2019b). Palp as in Figs 59B, 69B.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona melanosticta.

Clubiona zhangyongjingi Li & Blick, 2019

Clubiona transversa Zhang & Yin, 1998: 14, figs 16–19 (♀ only).

Clubiona zhangyongjingi Li & Blick, 2019: 131 (replacement name for C. transversa; ♂ mismatched).

Material examined

None.

Diagnosis and description

See Zhang and Yin (1998).

Distribution

China (Yunnan).

Remarks

Based on the original figures, the female is almost the same as that of C. melanosticta, and the male belongs to the C. ternatensis group and resembles C. kuu and C. subkuu.

Clubiona trivialis group

Microclubiona: Lohmander, 1944: 20 (type species C. trivialis C.L. Koch, 1834).

Clubiona trivialis group: Dondale and Redner 1976: 1155; Mikhailov 1995: 43.

Diagnosis

See Dondale and Redner (1976) and Yu and Li (2019a).

Description

See Dondale and Redner (1976) and Mikhailov (1995).

Composition and distribution

Currently, the trivialis group includes at least 28 species mainly distributed in Eurasia and Australia (Mikhailov 1995; Dondale and Redner 1982; Dong and Zhang 2016). Among these, at least 16 species (including the new species described here) have been recorded from China (see Table 6).

Comments

Lohmander (1944) established the genus Microclubiona with the type species C. trivialis. However, Lohmander’s work was neglected by North American taxonomists. Clubiona trivialis and related species were placed in Locket and Millidge’s (1951) Group III and Edwards’ (1958) Group I, respectively. The Clubiona trivialis group was formally established by Dondale and Redner (1976) and was redefined by Mikhailov (1995) based on 19 Holarctic species. The group had been separated from genus Clubiona sensu lato, to be resurrected to genus level by Wunderlich (2011) but latter synonymised with Clubiona by Mikhailov (2012). The present study follows Mikhailov (2012) and the WSC (2021) in regarding Microclubiona as a synonym of Clubiona rather than revalidating the generic status of the trivialis group. Consequently, we temporarily place the five Xishuangbanna species in Clubiona sensu lato and assign them to C. trivialis group.

Table 6.

Clubiona trivialis group.

Species name Known sex Distribution
1 C. amurensis Mikhailov, 1990 ♂♀ Russia (Far East), Japan (Hokkaido)
2 C. asrevida Ono, 1992 ♂♀ China (Taiwan)
3 C. baimaensis Song & Zhu, 1991 ♂♀ China (Hubei, Hunan, Sichuan)
4 C. basarukini Mikhailov, 1990 ♂♀ Russia (South Siberia, Far East), Mongolia, Japan (Hokkaido)
5 C. bicornis Yu & Li, 2019 ♂♀ China (Yunnan)
6 C. cheni Yu & Li, 2019 ♂♀ China (Yunnan)
7 C. diversa O. Pickard-Cambridge, 1862 ♂♀ Trans Palaearctic
8 C. duoconcava Zhang & Hu, 1991 ♂♀ South China
9 C. hedini Schenkel, 1936 China (Hunan, Gansu)
10 C. hooda Dong & Zhang, 2016 ♂♀ China (Hebei)
11 C. huaban Xin, Zhang, Li, Zeng & Yu, 2020 China (Guizhou)
12 C. insulana Ono, 1989 ♂♀ China (Taiwan), Japan (Ryukyu Is.)
13 C. janae Edwards, 1958 USA (California)
14 C. juvenis Simon, 1878 ♂♀ West Palaearctic
15 C. moesta Banks, 1896 ♂♀ Nearctic, China (Hunan, Hubei, Qinghai, Guizhou)
16 C. pygmaea Banks, 1892 ♂♀ Nearctic
17 C. quebecana Dondale & Redner, 1976 ♂♀ Nearctic
18 C. rostrata Paik, 1985 ♂♀ FE Palaearctic
19 C. subasrevida Yu & Li, 2019 ♂♀ China (Yunnan)
20 C. subquebecana Yu & Li, 2019 ♂♀ China (Yunnan)
21 C. subrostrata Zhang & Hu, 1991 ♂♀ China (Fujian, Hunan, Guizhou)
22 C. subtilis L. Koch, 1867 ♂♀ Trans Palaearctic
23 C. subtrivialis Strand, 1906 ♂♀ East Africa
24 C. subyangmingensis Gan & Wang, 2020 ♂♀ China (Guizhou)
25 C. transbaicalica Mikhailov, 1992 Baikal Lake
26 C. trivialis C. L. Koch, 1843 ♂♀ Holarctic
27 C. yangmingensis Hayashi & Yoshida, 1993 ♂♀ China (Taiwan)
28 C. menglun Yu & Li, sp. nov. China (Yunnan)

Key to C. trivialis group species occurring in Xishuangbanna (males)

1 Palp with prolateral tibial apophysis (Figs 61A, B, 71A, B) 2
Palp without prolateral tibial apophysis (Figs 61C, D, 71C, D) 3
2 Retrolateral tibial apophysis branched, both ventral and dorsal branches sharply pointed (Fig. 71A); embolar base bearing only dentiform process (Figs 61A, 71A) C. bicornis
Retrolateral tibial apophysis not branched, with a blunt tip (Fig. 71B); embolar base bearing two processes (Fig. 71B) C. cheni
3 Retrolateral tibial apophysis broad, flat and triangular, with sharp tip (Figs 61C, 71C) C. subasrevida
Retrolateral tibial apophysis small, thumb-like, with a blunt tip (Figs 61D, 71D) C. subquebecana

Key to C. trivialis group species occurring in Xishuangbanna (females)

1 Copulatory openings fused (Figs 78C, 86C) C. subasrevida
Copulatory openings separated (Figs 77F, 78A, B, D, 85F, 86A, B, D) 2
2 Spermathecae larger than bursae (Figs 93F, 94D) 3
Spermathecae smaller than bursae (Fig. 94A, B) 4
3 Spermathecae peanut-shaped (Fig. 93F) C. bicornis
Spermathecae subglobular (Fig. 94D) C. subquebecana
4 Spermathecae ellipsoidal (Fig. 94A) C. cheni
Spermathecae globular (Fig. 94B) C. menglun sp. nov.

Clubiona bicornis Yu & Li, 2019

Figs 48, 61A, 71A, 77F, 85F, 93F

Clubiona bicornis Yu & Li, 2019b: 221, figs 15A–E, 16A–C (♂).

Material examined

Type. Holotype ♂ (IZCAS Ar 34716), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, primary tropical seasonal rainforest, 21°55.035'N, 101°16.500'E, ca. 558 m, 22.VII.2007, G. Zheng leg. Other material examined. 1♂ (YHCLU0180), Jinghong City: Menga Town: Wengnan Village: secondary forest, 22°4.598'N, 100°22.134'E, ca. 1137 m, 30.VII.2012, Q.Y. Zhao and Z.G. Chen leg; 1♀ (YHCLU0099), XTBG, 48th km landmark in Menglun Nature Reserve, 21°58.704'N, 101°19.748'E, ca. 1088 m, 11.VIII.2011, G. Zheng et al. leg.

Diagnosis

Females of C. bicornis can be easily distinguished from other members of the group except C. amurensis (Mikhailov, 1990: 148, figs 21, 22) by the copulatory openings separated by one diameter (Figs 48A–C, 77F, 85F) (vs. copulatory openings fused or close together in almost all species of the C. trivialis group, including C. subasrevida and C. menglun sp. nov.; Figs 49A–C, 78B, C, 86B, C) and the lengthwise spermathecae (Figs 48D, E, 93F) (vs. spermathecae nearly globular in other species of the trivialis group, such as C. cheni and C. menglun sp. nov.; Figs 49D, E, 94A, B) but differ from the latter by the: (1) copulatory opening a small pore (Figs 48A–C, 77F, 85F) (vs. slit like in C. amurensis; Mikhailov, 1990: fig. 21); (2) peanut-shaped spermathecae (Figs 48D, E, 93F) (vs. elliptical in C. amurensis; Mikhailov, 1990: fig. 22); (3) proximal half of the copulatory ducts close together (Figs 48D, E, 93F) (vs. widely separated by more than four diameters in C. amurensis; Mikhailov, 1990: fig. 22); (4)fovea indistinct (Fig. 48F) (vs. fovea distinct in C. amurensis).

Description

Male. See Yu and Li (2019b). palp as in Figs 61A, 71A.

Female. (Fig. 48F, G): Total length 2.33; carapace 0.99 long, 0.77 wide; opisthosoma 1.34 long, 0.83 wide. Carapace, in profile almost flat, brown, slightly darker in front, with a pair of indistinct short lines running longitudinally from behind AME, fovea indistinct; ocular region slightly narrowed, cervical groove and radial grooves indistinct; tegument smooth, clothed with numerous short, fine setae. Eyes: AER almost straight, PER slightly recurved and slightly wider than AER in dorsal view. Eye sizes and interdistances: AME 0.05, ALE 0.06, PME 0.04, PLE 0.05, AMEAME 0.03, AMEALE 0.13, PMEPME 0.27, PMEPLE 0.08, MOQL 0.17, MOQA 0.13, MOQP 0.32. Chelicerae protruding and robust, coloured as carapace, with distinct lateral bulge, cheliceral furrow with three anterior and two posterior teeth Sternum pale brown, 0.64 long, 0.45 wide. Labium and endites coloured as carapace. Legs light brown, without distinct markings. Leg measurements: I 1.72 (0.48, 0.77, 0.35, 0.13), II 1.69 (0.51, 0.62, 0.40, 0.16), III 1.33 (0.41, 0.48, 0.28, 0.15), IV 1.82 (0.52, 0.78, 0.32, 0.20). Abdomen oval, cream coloured, numerous large pigmented markings prominently visible through integument except anteriorly and on the spinnerets.

Epigyne (Figs 48A–E, 77F, 85F, 93F). Epigynal plate slightly wider than long, the arrangement of the various parts of the vulva are indistinctly visible through the tegument. Copulatory openings indistinct, separated by one diameter, situated at medial portion of epigynal plate posterior margin. Hyaline copulatory ducts ascending in parallel, the proximal half close together, the distal half widely separated. Spermathecae close together, peanut- or gourd-shaped, ca. 2 × longer than wide. Fertilisation ducts curved and acicular, relatively long, > 1/3 spermathecae length, located on anterior surface of spermathecae. Bursae oblong, ca. 1.8 × longer than wide, with a smooth hyaline surface.

Distribution

Known only from Xishuangbanna.

Remarks

The female of the species is described for the first time.

Clubiona cheni Yu & Li, 2019

Figs 61B, 71B, 78A, 86A, 94A

Clubiona cheni Yu & Li, 2019a: 171, figs 13A–E, 14A–H (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34625), 1♀ (paratype, IZCAS Ar 34626), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, low evergreen forest, 21°53.794'N, 101°17.152'E, ca. 594 m, 27.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♂ (YHCLU0033), XTBG, secondary tropical forest, 21°54.168'N, 101°16.866'E, ca. 610 m, 31.VII.2018, Z.G. Chen leg; 1♀ (YHCLU0032), XTBG, low evergreen forest, 21°53.823'N, 101°17.072'E, ca. 613 m, 22.VII.2018, H. Yu et al. leg.

Diagnosis and description

See Yu and Li (2019a). Male palp as in Figs 61B, 71B, epigyne as in Figs 78A, 86A, 94A.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona bicornis.

Clubiona menglun Yu & Li, sp. nov.

Figs 49, 78B, 86B, 94B

Holotype

♀ (IZCAS-Ar 34762), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary tropical seasonal moist forest, 22°54.390'N, 101°16.811'E, ca. 612 m, 10.VIII.2007, G. Zheng leg. Other material examined. 1♀ (YHCLU0097), Jinghong City: Menga Town: Wengnan Village: secondary forest, 22°4.997'N, 100°22.223'E, ca. 1137 m, 25.VII.2012, Q.Y. Zhao and Z.G. Chen leg.

Etymology

The species name is derived from the name of the type locality; noun in apposition.

Diagnosis

Females of C. menglun sp. nov. resemble those of C. cheni (Figs 78A, 86A, 94A) by having a similar general shape of the vulva but can be separated from them by the close spermathecae (vs. separated by ca. 0.5 diameter) (cf. Fig. 94B and 94A) and by the globular bursae (vs. ellipsoidal) (cf. Fig. 94B and 94A).

Description

Female. Holotype (Fig. 49F, G): Total length 2.99; carapace 1.07 long, 0.82 wide; opisthosoma 1.83 long, 1.07 wide. Carapace orange, slightly darker in front, without distinct pattern, fovea almost indistinguishable; ocular region slightly narrowed, cervical groove indistinct; tegument smooth, clothed with numerous short, fine setae. Eyes: in dorsal view, both anterior and posterior eye rows recurved, PER slightly wider than AER. Eye sizes and interdistances: AME 0.06, ALE 0.03, PME 0.07, PLE 0.05, AMEAME 0.04, AMEALE 0.04, PMEPME 0.20, PMEPLE 0.04, MOQL 0.13, MOQA 0.12, MOQP 0.31. Chelicerae robust and brownish red, with conspicuous condyle, three promarginal and two retromarginal teeth. Sternum pale yellow, 0.67 long, 0.42 wide. Labium and endites coloured as chelicerae. Legs light yellowish white, without distinct markings. Leg measurements: I 1.77 (0.52, 0.70, 0.34, 0.21), II 2.02 (0.56, 0.86, 0.38, 0.22), III 1.50 (0.48, 0.53, 0.32, 0.17), IV 2.03 (0.60, 0.74, 0.46, 0.23). Abdomen oval, cream coloured, slightly darker dorsally, without pattern.

Epigyne (Figs 49A–E, 78B, 86B, 94B). Epigynal plate slightly longer than wide, vulva clearly visible through the tegument. Copulatory openings distinct, close together, located close to posterior margin of epigynal plate. Hyaline copulatory ducts long and slender, almost parallel, ascending dorsally, then ascending obliquely, finally entering the connecting piece between the spermathecae and bursae. Both spermathecae and bursae globular and smooth, the former anteriad and smaller than the latter. Spermathecae close together, and bursae separated by one diameter. Fertilisation ducts acicular, relatively long, more than half spermathecae diameter, on dorsal surfaces of spermathecae.

Male. Unknown.

Comments

Only two trivialis group species are known from males (Table 6): C. transbaicalica from South Siberia and C. huaban from Guizhou in China. We cannot rule out the possibility that C. transbaicalica is conspecific to C. menglun sp. nov. However, the probability is very small because of: (1) the long distance between the two type localities (Xishuangbanna is 3500 Km from the Selenga Distr.); (2) their different sizes (C. menglun sp. nov. is less than 3 mm, C. transbaicalica is 5.3–6.1 mm). C. menglun sp. nov. and C. huaban are considered separate species due to their different sizes (C. menglun sp. nov. with 3 mm vs. C. huaban with 4.7 mm; different colours (carapace orange, cream-coloured abdomen without pattern in C. menglun sp. nov. (vs. carapace light brown, yellowish brown abdomen marked with numerous brown spots in C. huaban).

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona subasrevida Yu & Li, 2019

Figs 61C, 71C, 78C, 86C, 94C

Clubiona subasrevida Yu & Li, 2019b: 221, figs 17A–E, 18A–H (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34717), 1♀ (paratype, IZCAS Ar 34718), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary tropical montane evergreen broad-leaved forest, 21°57.809'N, 101°12.173'E, ca. 888 m, 4.VIII.2007, G. Zheng leg. Other material examined. 1♂ (YHCLU0100), Huigang Village, monsoon forest, 21°37.027'N, 101°35.161'E, ca. 764 m, 12.VII.2012, Q.Y. Zhao and C.X. Gao leg; 1♀ (YHCLU0101), XTBG, 48th km landmark in Menglun Nature Reserve, 21°58.704'N, 101°19.748'E, ca. 1088 m, 12.VIII.2011, G. Zheng et al. leg.

Diagnosis and description

See Yu and Li (2019b). Male palp as in Figs 61C, 71C, epigyne as in Figs 78C, 86C, 94C.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona asrevida.

Clubiona subquebecana Yu & Li, 2019

Figs 61D, 71D, 78D, 86D, 94D

Clubiona subquebecana Yu & Li, 2019a: 174, figs 15A–E, 16A–H (♂♀).

Material examined

Types. Holotype ♂ (IZCAS Ar 34685), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary forest, 21°54.459'N, 101°16.755'E, ca. 644 m, 20.XI.2009, G. Tang and Z.Y. Yao leg; 1♀ (paratype, IZCAS Ar 34687), XTBG, G213 roadside, Anogeissus acuminata plantation, 21°53.819'N, 101°17.075'E, ca. 609 m, 27.XI.2009, G. Tang and Z.Y. Yao leg. Other material examined. 1♀ (YHCLU0103), Xiaolongha Village, 21°24.198'N, 101°37.013'E, ca. 801 m, 30.VI.2012, Q.Y. Zhao and C.X. Gao leg.

Diagnosis and description

See Yu and Li (2019a). Male palp as in Figs 61D, 71D, epigyne as in Figs 78D, 86D, 94D.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona quebecana.

Species not currently assigned to any group

Clubiona jiandan Yu & Li, 2019

Figs 62C, 72C, 80B, 88B, 96B

Clubiona jiandan Yu & Li, 2019b: 226, figs 19A–E, 20A–H (♂♀).

Material examined

Holotype ♂ (IZCAS Ar 34720), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, secondary tropical montane evergreen broad-leaved forest, 21°57.784'N, 101°11.947'E, ca. 895 m, 6.VIII.2007, G. Zheng leg; 1♀ (paratype, IZCAS Ar 34723), XTBG, secondary tropical montane evergreen broad-leaved forest, 21°57.528'N, 101°12.384'E, ca. 890 m, 6.VIII.2007, G. Zheng leg. Other material examined. 1♂ (YHCLU0066) and 1♀ (YHCLU0067), Xiaolongha Village, 21°24.198'N, 101°37.013'E, ca. 801 m, 30.VI.2012, Q.Y. Zhao and C.X. Gao leg.

Diagnosis and description

See Yu and Li (2019b). Male palp as in Figs 62C, 72C, epigyne as in Figs 80B, 88B, 96B.

Distribution

Known only from Xishuangbanna.

Most similar species

Clubiona yaoi.

Clubiona shuangsi Yu & Li, sp. nov.

Figs 50, 51, 62D, 72D, 80D, 88D, 96D

Holotype

♂ (IZCAS-Ar 34763, YHCLU0135), China: Yunnan Province: Xishuangbanna: Mengla County: Nanshahe Village: monsoon forest, 21°36.200'N, 101°34.384'E, ca. 826 m, 14.VII.2012, Q.Y. Zhao and C.X. Gao leg. Paratype: 1♀ (IZCAS-Ar 34764), same data as holotype. Other material examined. 1♀ (YHCLU0085), same data as holotype.

Etymology

The specific name is derived from the Chinese pinyin shuāng sī, which means ‘two filaments’, referring to the filiform embolus and conductor; noun in apposition.

Diagnosis

Males of C. shuangsi sp. nov. resemble those of C. biembolata (Deeleman-Reinhold 2001: 132, figs 67–69) in having a similar filiform embolus and conductor but differ by the retrolateral tibial apophysis with a relatively shorter tip, blunt, without inner apophysis (Figs 50B, 51E, 62D, 72D) (vs. tibial apophysis with long, acuminate tip, accompanied by a small inner apophysis; Deeleman-Reinhold 2001: figs 67, 68); and both the embolus and conductor shorter than the tegulum width (Figs 50C–E, 62D) (vs. embolus and conductor longer than tegulum width; Deeleman-Reinhold 2001: fig. 69). Females also resemble those of C. biembolata in having a small and rebordered atrium but can be recognised by the atrium-shaped nearly like an inverted triangle (Figs 51A, B, 80D, 88D) (vs. round; Deeleman-Reinhold 2001: fig. 70) and the bean-shaped spermathecae smaller than bursae (Figs 51C, D, 96D) (vs. tubular spermathecae with convoluted distal part, larger than bursae; Deeleman-Reinhold 2001: fig. 71).

Remarks

C. shuangsi sp. nov. resembles C. biembolata which was first described and assigned to the C. japonica group (called C. filicata group in the present paper) by Deeleman-Reinhold (2001) because of the characteristic copulatory organs (for a detailed diagnosis, see above). However, both species lack the dark pattern found on the dorsum of the opisthosoma in all existing members of the filicata group. Thus, there remains considerable uncertainty about placing the two species in the filicata group.

Description

Male. Holotype (Fig. 51F, G): Carapace 1.90 long, 1.36 wide. Carapace uniformly brown, without distinct pattern; ocular area distinctly narrowed, cervical groove and radial grooves indistinctly visible; tegument smooth, marginally clothed with short setae. Eyes: AER slightly recurved, PER slightly wider than AER, almost straight in dorsal view. Eye sizes and interdistances: AME 0.09, ALE 0.14, PME 0.11, PLE 0.09, AMEAME 0.07, AMEALE 0.04, PMEPME 0.17, PMEPLE 0.09, MOQL 0.29, MOQA 0.25, MOQP 0.42. Chelicerae robust and brown, with four promarginal and three retromarginal teeth. Sternum pale brown, 1.07 long, 0.69 wide. Labium and endites coloured as carapace. Legs light brown, without distinct markings. Leg measurements: I missing, II 4.81 (1.46, 1.92, 0.98, 0.46), III 3.86 (1.18, 1.25, 1.04, 0.42), IV 5.53 (1.53, 1.87, 1.63, 0.50). Abdomen missing.

Palp (Figs 50A–E, 51E, 62D, 72D). Tibia short, ca. 2 × shorter than cymbium, bearing group of ventral setae; RTA large, ca. as long as tibia, proximally broad and heavily sclerotised, distally thinner and partly membranous, tip blunt. Bulb oval, 1.9 × longer than wide; sperm duct distinct and sinuate, forming a double loop. Embolus filiform, arising at ca. 11 o’clock position, broad at base, gradually tapering toward apex, embolar tip pointing distally. Conductor originating from anterior membranous portion of tegulum, consisting of broad base and filiform distal part, base partly membranous and covering embolar apex, apex sharp and pointing prolatero-proximally.

Female. Paratype (Fig. 51H, I). Total length 4.07; carapace 1.82 long, 1.37 wide; opisthosoma 2.25 long, 1.40 wide. Eye sizes and interdistances: AME 0.10, ALE 0.13, PME 0.10, PLE 0.11, AMEAME 0.08, AMEALE 0.07, PMEPME 0.23, PMEPLE 0.15, MOQL 0.26, MOQA 0.23, MOQP 0.44. Chelicerae with three promarginal and two retromarginal teeth. Sternum 0.99 long, 0.67 wide. Leg measurements: I 3.60 (1.34, 1.28, 0.66, 0.42), II 3.91 (1.18, 1.57, 0.72, 0.45), III 3.24 (1.03, 1.08, 0.76, 0.37), IV 4.83 (1.34, 1.68, 1.38, 0.44). Colouration lighter than in male. Other characters as in male.

Epigyne (Figs 51A–D, 80D, 88D, 96D). Epigynal plate ca. 1.2 × wider than long, through which spermathecae and bursae are indistinctly apparent. Atrium small and shaped like an inverted triangle, with distinctly rebordered margin, ca. 1/3 epigyne length and width, anterior margin almost straight, posterior margin V-shaped. Copulatory openings located at anterolateral atrial borders. Copulatory ducts absent. Spermathecae bean-shaped, ca. 1.3 × longer than wide, separated by one diameter. Bursae ovoid, ca. 1.4 × longer than wide, close together, surface membranous and translucent, inside pigmented.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Clubiona wangchengi Yu & Li, sp. nov.

Figs 52, 80C, 88C, 96C

Holotype

♀ (IZCAS-Ar 34765), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, 48th km landmark in Menglun Nature Reserve, 21°58.704'N, 101°19.748'E, ca. 1088 m, 12.VIII.2011, G. Zheng et al. leg. Other material examined. 1♀ (YHCLU0087), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: Bubang Village, Parashorea cathayensis forest, 21°35.011'N, 101°35.013'E, ca. 680 m, 28.VII.2016, G. Zheng leg.

Etymology

This species is named after Mr. Cheng Wang (Tongren City, China) who has helped us greatly with this research; noun (name) in genitive case.

Diagnosis

Clubiona wangchengi sp. nov. resembles C. subkuu by the similar habitus: wide head, not much narrower than the carapace, and yellowish body (Fig. 52F, G; Yu and Li 2019a: 164, fig. 10G, H) but is consistently separable by the epigyne. The epigynes of both species share the similarly globular spermathecae and bursae and short and curved copulatory ducts which ascend obliquely but differ in the following: (1) the epigynal plate without ridges in C. wangchengi sp. nov. (Figs 52A–C; 80C, 88C) (vs. with blade-shaped ridges in C. subkuu; Figs 77B, 85B); (2) bursae located on the lateral sides of the spermathecae in C. wangchengi sp. nov. (Figs 52D, E; 96C) (vs. bursae located posterior to the spermathecae in C. subkuu; Fig. 93B).

Remarks

Clubiona wangchengi sp. nov. resembles some members of the C. ternatensis group by the wide head and the general shape of the vulva but can be distinguished from these species by the absence of epigynal ridges. Because all C. ternatensis group species have epigynal ridges (or hoods, or folds), there remains considerable uncertainty about placing this new species in the ternatensis group. In addition, this new species resembles some species of Pteroneta Deeleman-Reinhold, 2001, which is most similar to the C. ternatensis group in genital morphology. This new species can be separated from all known members of the genus Pteroneta by its unpatterned yellow body (Pteroneta has a pale green body, ventrally with lazulite blue spots). Despite the similarity of the general shape of the vulva in C. wangchengi sp. nov. and species of the ternatensis group and the Pteroneta, it is currently impossible to discern any obvious derived features (i.e., epigynal ridges and pale green body) that could indicate placement in the Clubiona ternatensis group or the genus Pteroneta.

Description

Female. Holotype (Fig. 52F, G): Total length 8.25; carapace 3.41 long, 2.24 wide; opisthosoma 4.84 long, 2.89 wide. Carapace orange, slightly darker in front, without distinct pattern, cephalic region slightly narrowed, cervical groove and radial grooves indistinct; tegument smooth, anteriorly clothed with sparse setae. Eyes: AER slightly recurved, PER slightly wider than AER, almost straight in dorsal view. Eye sizes and interdistances: AME 0.17, ALE 0.15, PME 0.15, PLE 0.13, AMEAME 0.08, AMEALE 0.11, PMEPME 0.39, PMEPLE 0.21, MOQL 0.43, MOQA 0.42, MOQP 0.69. Chelicerae robust and brownish red, with three teeth on promargin and two on retromargin. Sternum yellowish white, 1.72 long, 1.03 wide. Labium and endites coloured as carapace. Legs light yellow, without distinct markings. Leg measurements: I 5.63 (1.64, 2.45, 0.97, 0.56), II 6.00 (1.76, 2.60, 1.06, 0.58), III – (–, 2.83, 2.36, –), IV – (1.73, 1.36, 0.46, –). Abdomen elongate, oval, uniformly cream coloured, dorsum with a narrow, heart-shaped mark and two pairs of conspicuous muscle depressions; venter medially with two longitudinal dotted lines.

Epigyne (Figs 52A–E, 80C, 88C, 96C). Epigynal plate ca. 1.5 × wider than long, with spermathecae, bursae and ducts prominent through tegument. Copulatory openings large, located on the window (or chitinous structure) which is at the postero-lateral portion of the epigynal plate. Copulatory ducts short and curved, connected to bursae midway between epigastric fold and anterior surface of the spermathecae. Both spermathecae and bursae are globular, the former larger than the latter. Spermathecae sclerotised, close together. Fertilisation ducts acicular, relatively long, nearly equal to spermathecae diameter, located on anterolateral surface of spermathecae. Bursae located on the lateral sides of the spermathecae, separated by ca. two diameters. Bursal surface membranous and wrinkled, inside pigmented and sclerotised.

Male. Unknown.

Distribution

Known only from the type locality, Xishuangbanna, Yunnan, China.

Figures

Figure 1. 

Clubiona cochlearis, epigyne (A–E) and female habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: A = atrium; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm (equal for A–E); 1 mm (equal for F, G).

Figure 2. 

Holotype female of Clubiona dengpao sp. nov., epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: A = atrium; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm (equal for A–E); 1 mm (equal for F, G).

Figure 3. 

Male palp of Clubiona kurosawai A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 4. 

Clubiona kurosawai, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F Lateral view G dorsal view H ventral view. Abbreviations: BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm (equal for A–D); 2 mm (equal for E, F, equal for G, H).

Figure 5. 

Male palp of Clubiona moralis A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; PPA = prolateral patellar apophysis; RPA = retrolateral patellar apophysis; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 6. 

Clubiona moralis, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: BS = bursa; CD = copulatory duct; CO = copulatory opening; SP = spermatheca. Scale bars: 0.2 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 7. 

Male palp of Clubiona multidentata. A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; PPA = prolateral patellar apophysis; RPA = retrolateral patellar apophysis; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 8. 

Clubiona multidentata, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 9. 

Male palp of Clubiona parconcinna A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; PPA = prolateral patellar apophysis; RPA = retrolateral patellar apophysis; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 10. 

Clubiona parconcinna, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: AAM = atrial anterior margin; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.2 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 11. 

Male palp of Clubiona pollicaris A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: E = embolus; EB = embolar base; FA = femoral apophysis; PFR = prolateral femoral ridge; PPA = prolateral patellar apophysis; TA = tegular apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 12. 

Clubiona pollicaris, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: BS = bursa; CO = copulatory opening; SP = spermatheca. Scale bars: 0.2 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 13. 

Male palp of Clubiona rama A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.2 mm (equal for A, B, equal for C–E).

Figure 14. 

Male habitus of Clubiona rama. A dorsal view B ventral view C lateral view. Scale bar: 2 mm (equal for A–C).

Figure 15. 

Holotype female of Clubiona subdidentata sp. nov., epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: AM = atrial membrane; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm (equal for A–E); 1 mm (equal for F, G).

Figure 16. 

Male palp of Clubiona submoralis A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; PPA = prolateral patellar apophysis; RPA = retrolateral patellar apophysis; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 17. 

Clubiona submoralis, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.2 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 18. 

Holotype female of Clubiona tixing sp. nov., epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: A = atrium; AM = atrial membrane; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm (equal for A–E); 1 mm (equal for F, G).

Figure 19. 

Clubiona tiane, epigyne (A–E) and female habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: A = atrium; BS = bursa; CO = copulatory opening; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.1 mm (equal for A–E); 1 mm (equal for F, G).

Figure 20. 

Male palp of the holotype of Clubiona xiaoci sp. nov. A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: E = embolus; EB = embolar base; RPA = retrolateral patellar apophysis; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 21. 

Clubiona xiaoci sp. nov., female paratype and male holotype, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: AAM = atrial anterior margin; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 22. 

Holotype female of Clubiona xiaokong sp. nov., epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: A = atrium; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm (equal for A–E); 2 mm (equal for F, G).

Figure 23. 

Holotype female of Clubiona yejiei sp. nov., epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: A = atrium; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm (equal for A–E); 1 mm (equal for F, G).

Figure 24. 

Holotype female of Clubiona zhaoi sp. nov., epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: A = atrium; BS = bursa; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.2 mm (equal for A–E); 1 mm (equal for F, G).

Figure 25. 

Male palp of the holotype of Clubiona zhigangi sp. nov. A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.2 mm (equal for A, B, equal for C–E).

Figure 26. 

Clubiona zhigangi sp. nov., female paratype and male holotype, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: A = atrium; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca; SH = spermathecal head. Scale bars: 0.2 mm (equal for A–D); 2 mm (equal for E, F, equal for G, H).

Figure 27. 

Holotype female of Clubiona mii sp. nov., epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view Fdorsal view G ventral view. Abbreviations: BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; R = epigynal ridge; SP = spermatheca. Scale bars: 0.1 mm (equal for A–E); 1 mm (equal for F, G).

Figure 28. 

Male palp of the holotype of Clubiona subtongi sp. nov. A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; TH = tegular hump. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 29. 

Habitus of the male holotype of Clubiona subtongi sp. nov. A dorsal view B ventral view C lateral view. Scale bar: 1 mm (equal for A–C).

Figure 30. 

Male palp of Clubiona abnormis A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; TA = tegular hump. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 31. 

Male habitus of Clubiona abnormis A dorsal view B ventral view C lateral view. Scale bar: 1 mm (equal for A–C).

Figure 32. 

Male palp of the holotype of Clubiona banna sp. nov. A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; TA = tegular hump. Scale bars: 0.2 mm (equal for A, B, equal for C–E).

Figure 33. 

Clubiona banna sp. nov., female paratype and male holotype, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F Lateral view G dorsal view H ventral view. Abbreviations: A = atrium; BS = bursa; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.2 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 34. 

Male palp of Clubiona circulata A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis. Scale bars: 0.2 mm (equal for A, B, equal for C–E).

Figure 35. 

Clubiona circulata, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.2 mm (equal for A–D); 2 mm (equal for E, F, equal for G, H).

Figure 36. 

Male palp of Clubiona reichlini A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis. Scale bars: 0.2 mm (equal for A, B, equal for C–E).

Figure 37. 

Clubiona reichlini, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: A = atrium; AAM = atrial anterior margin; BS = bursa; CO = copulatory opening; FD = fertilisation duct; SB = spermathecal base; SH = spermathecal head; SP = spermatheca; SS = spermathecal stalk. Scale bars: 0.1 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 38. 

Male palp of Clubiona filicata A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; TA = tegular hump. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 39. 

Male habitus of Clubiona filicata A dorsal view B ventral view C lateral view. Scale bar: 1 mm (equal for A–C).

Figure 40. 

Male palp of Clubiona grucollaris A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; TA = tegular hump. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 41. 

Clubiona grucollaris, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: A = atrium; AAM = atrial anterior margin; BS = bursa; CO = copulatory opening; FD = fertilisation duct; SB = spermathecal base; SH = spermathecal head; SP = spermatheca; SS = spermathecal stalk. Scale bars: 0.2 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 42. 

Male palp of Clubiona lala A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; TA = tegular hump. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 43. 

Clubiona lala, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: A = atrium; AAM = atrial anterior margin; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SB = spermathecal base; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm (equal for A–D); 2 mm (equal for E, F, equal for G, H).

Figure 44. 

Male palp of Clubiona melanosticta A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 45. 

Clubiona melanosticta, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: A = atrium; BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SB = spermathecal base; SH = spermathecal head; SP = spermatheca; SS = spermathecal stalk. Scale bars: 0.1 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 46. 

Male palp of Clubiona suthepica A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 47. 

Clubiona suthepica, epigyne (A–D), male habitus (E, F) and female habitus (G, H) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E dorsal view F lateral view G dorsal view H ventral view. Abbreviations: A = atrium; AAM = atrial anterior margin; BS = bursa; CD = copulatory duct; CO = copulatory opening; SB = spermathecal base; SH = spermathecal head; SP = spermatheca. Scale bars: 0.1 mm (equal for A–D); 1 mm (equal for E, F, equal for G, H).

Figure 48. 

Clubiona bicornis, epigyne (A–E) and female habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm (equal for A–E); 0.5 mm (equal for F, G).

Figure 49. 

Holotype female of Clubiona menglun sp. nov. epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view Fdorsal view G ventral view. Abbreviations: BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm (equal for A–E); 1 mm (equal for F, G).

Figure 50. 

Male palp of the holotype of Clubiona shuangsi sp. nov. A prolateral view B retrolateral view C bulb, prolateral view D bulb, ventral view E bulb, retrolateral view. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm (equal for A, B, equal for C–E).

Figure 51. 

Clubiona shuangsi sp. nov., female paratype and male holotype, epigyne (A–D), tibia of male palp (E), male habitus (F, G) and female habitus (H, I) A intact, ventral view B cleared, ventral view C cleared, dorsal view D cleared, dorsal view E ventral view F dorsal view G lateral view H dorsal view I ventral view. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; BS = bursa; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm (equal for A–D); 0.1 mm (E); 1 mm (equal for E, G, equal for H, I).

Figure 52. 

Holotype female of Clubiona wangchengi sp. nov., epigyne (A–E) and habitus (F, G) A intact, ventral view B cleared, ventral view C cleared, ventral view D cleared, dorsal view E cleared, dorsal view F dorsal view G ventral view. Abbreviations: BS = bursa; CD = copulatory duct; CO = copulatory opening; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm (equal for A–E); 1 mm (equal for F, G).

Figure 53. 

Clubiona spp. of the C. corticalis group, male palp, ventral view A C. cochlearis B C. rama C C. subrama D C. zhigangi sp. nov., holotype. Abbreviations: C = conductor; E = embolus. Scale bars: 0.2 mm.

Figure 54. 

Clubiona spp. of the C. corticalis group, male palp, ventral view A C. didentata B C. kai C C. subyaginumai D C. tiane. Abbreviations: C = conductor; E = embolus; EB = embolar base; TA = tegular apophysis. Scale bars: 0.1 mm.

Figure 55. 

Clubiona spp. of the C. corticalis group, male palp, ventral view A C. moralis B C. submoralis C C. parconcinna D C. xiaoci sp. nov., holotype. Abbreviations: C = conductor; E = embolus; PPA = prolateral patellar apophysis; RPA = retrolateral patellar apophysis. Scale bars: 0.1 mm.

Figure 56. 

Clubiona spp. of the C. corticalis group, male palp, ventral view A C. kurosawai B C. multidentata C C. pollicaris. Abbreviations: C = conductor; E = embolus; EB = embolar base; FA = femoral apophysis; PPA = prolateral patellar apophysis; RPA = retrolateral patellar apophysis; PFR = prolateral femoral ridge; TA = tegular apophysis; VTA, ventral tibial apophysis. Scale bars: 0.1 mm.

Figure 57. 

Clubiona spp. of the C. ternatensis group, male palp, ventral view A C. theoblicki B C. tongi C C. subkuu D C. subtongi sp. nov., holotype E C. zhengi. Abbreviations: E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; TH = tegular hump. Scale bars: 0.1 mm.

Figure 58. 

Clubiona spp. of the C. filicata group, male palp, ventral view A C. reichlini B C. filicata C C. banna sp. nov., holotype. Abbreviations: C = conductor; E = embolus; EB = embolar base; TA = tegular apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 59. 

Clubiona spp. of the C. filicata group, male palp, ventral view A C. melanosticta B C. zhanggureni C C. circulata D C. suthepica. Abbreviations: C = conductor; E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 60. 

Clubiona spp. of the C. filicata group, male palp, ventral view A C. grucollaris B C. lala C C. abnormis D C. yueya. Abbreviations: C = conductor; E = embolus; EB = embolar base; TA = tegular apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 61. 

Clubiona spp. of the C. trivialis group, male palp, ventral view A C. bicornis B C. cheni C C. subasrevida D C. subquebecana. Abbreviations: C = conductor; E = embolus; EB = embolar base; TH = tegular hump; PTA = prolateral tibial apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 62. 

Clubiona spp., male palp, ventral view (A, C, D) and dorsal view (B) A, B C. yaoi C C. jiandan D C. shuangsi sp. nov., holotype. Abbreviations: C = conductor; DCA = dorsal cymbial apophysis; E = embolus; EB = embolar base; PTA = prolateral tibial apophysis; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm.

Figure 63. 

Clubiona spp. of the C. corticalis group, male palp, retrolateral view A C. cochlearis B C. rama C C. subrama D C. zhigangi sp. nov., holotype. Abbreviations: C = conductor; E = embolus; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.2 mm.

Figure 64. 

Clubiona spp. of the C. corticalis group, male palp, retrolateral view A C. didentata B C. kai C C. subyaginumai D C. tiane. Abbreviations: C = conductor; E = embolus; TA = tegular apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 65. 

Clubiona spp. of the C. corticalis group, male palp, retrolateral view A C. moralis B C. submoralis C C. parconcinna D C. xiaoci sp. nov., holotype. Abbreviations: C = conductor; E = embolus; RPA = retrolateral patellar apophysis; RTA, retrolateral tibial apophysis; VTA, ventral tibial apophysis. Scale bars: 0.1 mm.

Figure 66. 

Clubiona spp. of the C. corticalis group, male palp, retrolateral view A C. kurosawai B C. multidentata C C. pollicaris. Abbreviations: E = embolus; FA = femoral apophysis; PFR = prolateral femoral ridge; RPA = retrolateral patellar apophysis; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm.

Figure 67. 

Clubiona spp. of the C. ternatensis group, male palp, retrolateral view A C. theoblicki B C. tongi C C. subkuu D C. subtongi sp. nov., holotype E C. zhengi. Abbreviations: E = embolus; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm.

Figure 68. 

Clubiona spp. of the C. filicata group, male palp, retrolateral view A C. reichlini B C. filicata C C. banna sp. nov., holotype. Abbreviations: C = conductor; E = embolus; TA = tegular apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 69. 

Clubiona spp. of the C. filicata group, male palp, retrolateral view A C. melanosticta B C. zhanggureni C C. circulata D C. suthepica. Abbreviations: C = conductor; E = embolus; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 70. 

Clubiona spp. of the C. filicata group, male palp, retrolateral view A C. grucollaris B C. lala C C. abnormis D C. yueya. Abbreviations: C = conductor; E = embolus; TA = tegular apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 71. 

Clubiona spp. of the C. trivialis group, male palp, retrolateral view A C. bicornis B C. cheni C C. subasrevida D C. subquebecana. Abbreviations: C = conductor; E = embolus; PTA = prolateral tibial apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 72. 

Clubiona spp., male palp, prolateral view (A) and retrolateral view (B–D) A, B C. yaoi C C. jiandan D C. shuangsi sp. nov., holotype. Abbreviations: C = conductor; DCA = dorsal cymbial apophysis; E = embolus; EB = embolar base; PTA = prolateral tibial apophysis; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm.

Figure 73. 

Clubiona spp. of the C. corticalis group, epigyne, intact, ventral view A C. cochlearis B C. dengpao sp. nov., holotype C C. yejiei sp. nov., holotype D C. tixing sp. nov., holotype E C. subrama F C. zhigangi sp. nov., paratype. Abbreviations: A = atrium; AM = atrial membrane; CO = copulatory opening. Scale bars: 0.2 mm.

Figure 74. 

Clubiona spp. of the C. corticalis group, epigyne, intact, ventral view A C. xiaokong sp. nov., holotype B C. zhaoi sp. nov., holotype C C. kai D C. tiane E C. didentata F C. subdidentata sp. nov., holotype. Abbreviations: A = atrium; AM = atrial membrane; CO = copulatory opening. Scale bars: 0.2 mm.

Figure 75. 

Clubiona spp. of the C. corticalis group, epigyne, intact, ventral view A C. moralis B C. submoralis C C. parconcinna D C. multidentata E C. xiaoci sp. nov., paratype F C. subyaginumai. Abbreviations: AAM = atrial anterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 76. 

Clubiona spp. of the C. corticalis group, epigyne, intact, ventral view A C. kurosawai B C. pollicaris. Abbreviations: CO = copulatory opening. Scale bars: 0.2 mm.

Figure 77. 

Clubiona spp. of the C. ternatensis group (A–E) and the C. trivialis group (F), epigyne, intact, ventral view A C. mii sp. nov. holotype B C. subkuu C C. theoblicki D C. tongi E C. zhengi F C. bicornis. Abbreviations: A = atrium; CO = copulatory opening; R = epigynal ridge. Scale bars: 0.1 mm.

Figure 78. 

Clubiona spp. of the C. trivialis group (A–D) and the C. filicata group (E, F), epigyne, intact, ventral view A C. cheni B C. menglun sp. nov., holotype C C. subasrevida D C. subquebecana E C. banna sp. nov., paratype F C. melanosticta. Abbreviations: A = atrium; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 79. 

Clubiona spp. of the C. filicata group, epigyne, intact, ventral view A C. circulata B C. reichlini C C. grucollaris D C. lala E C. suthepica F C. yueya. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 80. 

Clubiona spp., epigyne, intact, ventral view A C. yaoi B C. jiandan C C. wangchengi sp. nov., holotype D C. shuangsi sp. nov., paratype. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 81. 

Clubiona spp. of the C. corticalis group, epigyne, cleared, ventral view A C. cochlearis B C. dengpao sp. nov., holotype C C. yejiei sp. nov., holotype D C. tixing sp. nov., holotype E C. subrama F C. zhigangi sp. nov., paratype. Abbreviations: A = atrium; AM = atrial membrane; CO = copulatory opening. Scale bars: 0.2 mm.

Figure 82. 

Clubiona spp. of the C. corticalis group, epigyne, cleared, ventral view A C. xiaokong sp. nov., holotype B C. zhaoi sp. nov., holotype C C. kai D C. tiane E C. didentata F C. subdidentata sp. nov., holotype. Abbreviations: A = atrium; AM = atrial membrane; CO = copulatory opening. Scale bars: 0.2 mm.

Figure 83. 

Clubiona spp. of the C. corticalis group, epigyne, cleared, ventral view A C. moralis B C. submoralis C C. parconcinna D C. multidentata E C. xiaoci sp. nov., paratype F C. subyaginumai. Abbreviations: AAM = atrial anterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 84. 

Clubiona spp. of the C. corticalis group, epigyne, cleared, ventral view A C. kurosawai B C. pollicaris. Abbreviations: CO = copulatory opening. Scale bars: 0.2 mm.

Figure 85. 

Clubiona spp. of the C. ternatensis group (A–E) and the C. trivialis group (F), epigyne, cleared, ventral view A C. mii sp. nov., holotype B C. subkuu C C. theoblicki D C. tongi E C. zhengi F C. bicornis. Abbreviations: A = atrium; CO = copulatory opening; R = epigynal ridge. Scale bars: 0.1 mm.

Figure 86. 

Clubiona spp. of the C. trivialis group (A–D) and the C. filicata group (E, F), epigyne, cleared, ventral view A C. cheni B C. menglun sp. nov., holotype C C. subasrevida D C. subquebecana E C. banna sp. nov., paratype F C. melanosticta. Abbreviations: A = atrium; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 87. 

Clubiona spp. of the C. filicata group, epigyne, cleared, ventral view A C. circulata B C. reichlini C C. grucollaris D C. lala E C. suthepica F C. yueya. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 88. 

Clubiona spp., epigyne, cleared, ventral view A C. yaoi B C. jiandan C C. wangchengi sp. nov., holotype D C. shuangsi sp. nov., paratype. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 89. 

Clubiona spp. of the C. corticalis group, vulva, cleared dorsal view A C. cochlearis B C. dengpao sp. nov., holotype C C. yejiei sp. nov., holotype D C. tixing sp. nov., holotype E C. subrama F C. zhigangi sp. nov., paratype. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm.

Figure 90. 

Clubiona spp. of the C. corticalis group, vulva, cleared, dorsal view A C. xiaokong sp. nov., holotype B C. zhaoi sp. nov., holotype C C. kai D C. tiane E C. didentata F C. subdidentata sp. nov., holotype. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm.

Figure 91. 

Clubiona spp. of the C. corticalis group, vulva, cleared, dorsal view A C. moralis B C. submoralis C C. parconcinna D C. multidentata E C. xiaoci sp. nov., paratype F C. subyaginumai. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm.

Figure 92. 

Clubiona spp. of the C. corticalis group, vulva, cleared, dorsal view A C. kurosawai B C. pollicaris. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm.

Figure 93. 

Clubiona spp. of the C. ternatensis group (A–E) and the C. trivialis group (F), vulva, cleared, dorsal view A C. mii sp. nov., holotype B C. subkuu C C. theoblicki D C. tongi E C. zhengi F C. bicornis. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm.

Figure 94. 

Clubiona spp. of the C. trivialis group (A–D) and the C. filicata group (E, F), vulva, cleared, dorsal view A C. cheni B C. menglun sp. nov., holotype C C. subasrevida D C. subquebecana E C. banna sp. nov., paratype F C. melanosticta. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SB = spermathecal base; SH = spermathecal head; SP = spermatheca; SS = spermathecal stalk. Scale bars: 0.1 mm.

Figure 95. 

Clubiona spp. of the C. filicata group, vulva, cleared, dorsal view A C. circulata B C. reichlini C C. grucollaris D C. lala E C. suthepica F C. yueya. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SB = spermathecal base; SH = spermathecal head; SP = spermatheca; SS = spermathecal stalk. Scale bars: 0.1 mm.

Figure 96. 

Clubiona spp., vulva, cleared, dorsal view A C. yaoi B C. jiandan C C. wangchengi sp. nov., holotype D C. shuangsi sp. nov., female paratype. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm.

Acknowledgements

We thank Yuri M. Marusik (Magadan, Russia), Mikhail M. Omelko (Vladivostok, Russia), and Feng Zhang (Baoding, China) for providing constructive comments on the manuscript. We thank Zhiyuan Yao (Shenyang, China) and Yucheng Lin (Chengdu, China) for providing constructive comments on an earlier version of the illustrations. Sarah Crews (San Francisco, USA) checked the English of final draft. Theo Blick (Hummeltal, Germany) checked the etymologies. Yanfeng Tong (Shenyang, China), Zhigang Chen (Beijing, China), Zilong Bai (Beijing, China), Yejie Lin (Beijing, China), Shijia Liu (Shenyang, China), Cheng Wang (Tongren, China), Xiaoqi Mi (Tongren, China), Jiahui Gan (Tongren, China), Yuanfa Yang (Tongren, China), and Hong Liu (Tongren, China) kindly helped in collecting the specimens. We thank Xiaoqing Zhang (Beijing, China), Fengyuan Li (Beijing, China), He Zhang (Hubei, China), Jiayuan Xin (Guiyang, China), and Da Wang (Guiyang, China) for help with molecular procedures. This study was supported by the National Natural Science Foundation of China to Hao Yu (NSFC-32060113/31702006), Jianshuang Zhang (NSFC-82060779), the Natural Science Foundation of Guizhou Province to Hao Yu ([2020]1Y081), PhD grant from Guizhou Normal University to Jianshuang Zhang (11904/0517069), Guizhou Education University Academic Discipline Project (2019YLPYXKB01), and Guizhou provincial first-class major (biological science) project (Education department of Guizhou Province [2019] 46).

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