Taxonomic studies on the sac spider genus Clubiona (Araneae, Clubionidae) from Xishuangbanna Rainforest, China

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.

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

Note: A total of 51 Clubiona species is reported from Xishuangbanna, of which 47 (marked with asterisks) were collected by the authors. The table includes only references that explicitly documented that the species was found in Xishuangbanna. For a complete list of taxonomic references, see taxonomy below and WSC (2021).

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
AME–AME	distance between AMEs
AME–ALE	distance between AME and ALE
PME–PME	distance between PMEs
PME–PLE	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

Genus. Clubiona

Latreille, 1804

96D05C06-E0E3-5A5D-B3CB-9B2608BC0B3B

• 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)

Figure 31.

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

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 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 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 39.

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

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 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 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 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 53.

Clubiona spp. of the C. corticalis group, male palp, ventral view AC. cochlearisBC. ramaCC. subramaDC. 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 AC. didentataBC. kaiCC. subyaginumaiDC. 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 AC. moralisBC. submoralisCC. parconcinnaDC. 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 AC. kurosawaiBC. multidentataCC. 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 63.

Clubiona spp. of the C. corticalis group, male palp, retrolateral view AC. cochlearisBC. ramaCC. subramaDC. 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 AC. didentataBC. kaiCC. subyaginumaiDC. 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 AC. moralisBC. submoralisCC. parconcinnaDC. 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 AC. kurosawaiBC. multidentataCC. 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 57.

Clubiona spp. of the C. ternatensis group, male palp, ventral view AC. theoblickiBC. tongiCC. subkuuDC. subtongi sp. nov., holotype EC. zhengi. Abbreviations: E = embolus; EB = embolar base; RTA = retrolateral tibial apophysis; TH = tegular hump. Scale bars: 0.1 mm.

Figure 62.

Clubiona spp., male palp, ventral view (A, C, D) and dorsal view (B) A, BC. yaoiCC. jiandanDC. 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 61.

Clubiona spp. of the C. trivialis group, male palp, ventral view AC. bicornisBC. cheniCC. subasrevidaDC. 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 72.

Clubiona spp., male palp, prolateral view (A) and retrolateral view (B–D) A, BC. yaoiCC. jiandanDC. 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 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 71.

Clubiona spp. of the C. trivialis group, male palp, retrolateral view AC. bicornisBC. cheniCC. subasrevidaDC. subquebecana. Abbreviations: C = conductor; E = embolus; PTA = prolateral tibial apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

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.

Figure 78.

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

Figure 81.

Clubiona spp. of the C. corticalis group, epigyne, cleared, ventral view AC. cochlearisBC. dengpao sp. nov., holotype CC. yejiei sp. nov., holotype DC. tixing sp. nov., holotype EC. subramaFC. 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 AC. xiaokong sp. nov., holotype BC. zhaoi sp. nov., holotype CC. kaiDC. tianeEC. didentataFC. 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 AC. moralisBC. submoralisCC. parconcinnaDC. multidentataEC. xiaoci sp. nov., paratype FC. 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 AC. kurosawaiBC. pollicaris. Abbreviations: CO = copulatory opening. Scale bars: 0.2 mm.

Figure 88.

Clubiona spp., epigyne, cleared, ventral view AC. yaoiBC. jiandanCC. wangchengi sp. nov., holotype DC. shuangsi sp. nov., paratype. 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 AC. yaoiBC. jiandanCC. wangchengi sp. nov., holotype DC. shuangsi sp. nov., paratype. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 77.

Clubiona spp. of the C. ternatensis group (A–E) and the C. trivialis group (F), epigyne, intact, ventral view AC. mii sp. nov. holotype BC. subkuuCC. theoblickiDC. tongiEC. zhengiFC. bicornis. Abbreviations: A = atrium; CO = copulatory opening; R = epigynal ridge. Scale bars: 0.1 mm.

Figure 85.

Clubiona spp. of the C. ternatensis group (A–E) and the C. trivialis group (F), epigyne, cleared, ventral view AC. mii sp. nov., holotype BC. subkuuCC. theoblickiDC. tongiEC. zhengiFC. bicornis. Abbreviations: A = atrium; CO = copulatory opening; R = epigynal ridge. Scale bars: 0.1 mm.

Figure 93.

Clubiona spp. of the C. ternatensis group (A–E) and the C. trivialis group (F), vulva, cleared, dorsal view AC. mii sp. nov., holotype BC. subkuuCC. theoblickiDC. tongiEC. zhengiFC. 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 AC. cheniBC. menglun sp. nov., holotype CC. subasrevidaDC. subquebecanaEC. banna sp. nov., paratype FC. 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 96.

Clubiona spp., vulva, cleared, dorsal view AC. yaoiBC. jiandanCC. wangchengi sp. nov., holotype DC. shuangsi sp. nov., female paratype. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm.

Clubiona milingae

group

07247417-2447-5EC1-8931-3B303D986027

• 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

0FA33ADC-8D7C-540F-BA42-050592D8C067

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

A875642A-381D-58DF-8958-D1CBDDC6A0E7

• 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.

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)

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).

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 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 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 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).

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.

Figure 76.

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

Figure 89.

Clubiona spp. of the C. corticalis group, vulva, cleared dorsal view AC. cochlearisBC. dengpao sp. nov., holotype CC. yejiei sp. nov., holotype DC. tixing sp. nov., holotype EC. subramaFC. 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 AC. xiaokong sp. nov., holotype BC. zhaoi sp. nov., holotype CC. kaiDC. tianeEC. didentataFC. subdidentata sp. nov., holotype. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SH = spermathecal head; SP = spermatheca. Scale bars: 0.2 mm.

Figure 92.

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

Figure 73.

Clubiona spp. of the C. corticalis group, epigyne, intact, ventral view AC. cochlearisBC. dengpao sp. nov., holotype CC. yejiei sp. nov., holotype DC. tixing sp. nov., holotype EC. subramaFC. 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 AC. xiaokong sp. nov., holotype BC. zhaoi sp. nov., holotype CC. kaiDC. tianeEC. didentataFC. 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 AC. moralisBC. submoralisCC. parconcinnaDC. multidentataEC. xiaoci sp. nov., paratype FC. subyaginumai. Abbreviations: AAM = atrial anterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 91.

Clubiona spp. of the C. corticalis group, vulva, cleared, dorsal view AC. moralisBC. submoralisCC. parconcinnaDC. multidentataEC. xiaoci sp. nov., paratype FC. subyaginumai. Abbreviations: BS = bursa; CD = copulatory duct; FD = fertilisation duct; SP = spermatheca. Scale bars: 0.1 mm.

Clubiona cochlearis

Yu & Li, 2019

17F5C4EF-C79C-5D1A-A3A9-A963C302289B

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

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).

• 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 bailloniiplantation, 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, AME–AME 0.21, AME–ALE 0.18, PME–PME 0.49, PME–PLE 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.

D8CFF7E2-DDA7-5FF6-989A-D3AA7DCA8792

http://zoobank.org/63A97FF6-8968-4584-BDBA-536500AA7479

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, AME–AME 0.18, AME–ALE 0.15, PME–PME 0.44, PME–PLE 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

234F835B-2AB2-59B5-93FF-F2D77D8116C3

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

1F0298CF-F5B6-5FD1-8710-2F5303FD4FE1

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 acuminataplantation, 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

BBCD68C7-3C1A-5396-BB5D-7C3130473D0A

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

Figure 3.

Male palp of Clubiona kurosawaiA 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).

• 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

99AC1803-DF52-511B-ADAD-4D46950A4065

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

Figure 5.

Male palp of Clubiona moralisA 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).

• 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 bailloniiplantation, 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 acuminataplantation, 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

D6507CB7-24B8-5946-B0E4-FACDD9B8B0B3

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

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).

• 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, 48^th 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

89335C6E-05DD-51F0-B7A1-97C4D89EC1E9

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

Figure 9.

Male palp of Clubiona parconcinnaA 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).

• 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, 55^th 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

123B10DD-D121-5576-AD9B-7BD005CA2148

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

Figure 11.

Male palp of Clubiona pollicarisA 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).

• 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

99CCB380-6207-5074-B15B-F741ACFE603F

Figs 13 , 14 , 53B , 63B

Figure 13.

Male palp of Clubiona ramaA 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).

• 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.

2FE4A8C4-00ED-5CA3-A665-AFE983608CB0

http://zoobank.org/B4B90700-E09E-430A-9B58-C80D6CF84084

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, AME–AME 0.06, AME–ALE 0.07, PME–PME 0.19, PME–PLE 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

00A116D3-C155-5682-AD1D-A255A514AA3F

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

Figure 16.

Male palp of Clubiona submoralisA 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).

• 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

ADC9EC07-E2D8-5ECF-A1EA-FB4A45B34127

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

7A96D7F7-2BF8-5826-B524-8B375F5E1B26

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.

31DC512F-7298-5C24-B97D-462BECB43C52

http://zoobank.org/0E0C7ACD-01A4-4D3F-833A-0F84AE4285AB

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, AME–AME 0.12, AME–ALE 0.13, PME–PME 0.39, PME–PLE 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

F51979B6-C35D-520B-8AE5-94513D150F01

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

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).

• 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, AME–AME 0.09, AME–ALE 0.06, PME–PME 0.17, PME–PLE 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.

BB1A2289-7B22-5ADB-BEC1-3485D33279D5

http://zoobank.org/1044ED46-1096-46CF-A856-CF991E21B404

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

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).

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, AME–AME 0.07, AME–ALE 0.05, PME–PME 0.10, PME–PLE 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, AME–AME 0.05, AME–ALE 0.05, PME–PME 0.19, PME–PLE 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.

AA593285-1B9F-5A28-9E05-1CBB1B8D4490

http://zoobank.org/D63C0094-D38F-4FFC-9937-D0A336324E28

Figs 22 , 74A , 82A , 90A

Holotype.

♀ (IZCAS-Ar 34753), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Anogeissus acuminataplantation (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, AME–AME 0.16, AME–ALE 0.11, PME–PME 0.35, PME–PLE 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.

EA3DD79B-0ED0-51BF-A944-31F7AF6A9355

http://zoobank.org/2EDB9A18-CA1C-4C1E-BA1B-3D83D01DC25A

Figs 23 , 73C , 81C , 89C

Holotype.

♀ (IZCAS-Ar 34754), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Paramichelia bailloniiplantation, 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, AME–AME 0.25, AME–ALE 0.85, PME–PME 0.45, PME–PLE 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.

6809F123-0C64-50E7-B3E1-C21BD9DE7F71

http://zoobank.org/DB382D68-B622-49C7-A799-773EDB02E05E

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, AME–AME 0.11, AME–ALE 0.08, PME–PME 0.32, PME–PLE 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.

D897AC49-B437-5C43-B346-4146C250E3FA

http://zoobank.org/03F6ECE2-ACAF-4DF3-9169-C04CF5BD84AF

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

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).

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 bailloniiplantation, 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, AME–AME 0.19, AME–ALE 0.13, PME–PME 0.30, PME–PLE 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, AME–AME 0.12, AME–ALE 0.11, PME–PME 0.35, PME–PLE 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

E0E32BC8-B6DE-555A-B263-E393AD91C451

• 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).

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)

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.

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.

Figure 67.

Clubiona spp. of the C. ternatensis group, male palp, retrolateral view AC. theoblickiBC. tongiCC. subkuuDC. subtongi sp. nov., holotype EC. zhengi. Abbreviations: E = embolus; RTA = retrolateral tibial apophysis; VTA = ventral tibial apophysis. Scale bars: 0.1 mm.

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

4DDD6DF5-CD3B-5BAF-B3E6-F056F8D33C1D

• 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.

862FA171-CCCC-5B61-AA6B-EE79FBFE1F51

http://zoobank.org/3C857D44-FF2D-4C7B-B459-4B83F0AB2361

Figs 27 , 77A , 85A , 93A

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).

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, AME–AME 0.04, AME–ALE 0.02, PME–PME 0.17, PME–PLE 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

C7CF5BC1-ECC2-5393-844E-44DD9FD75FA1

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.

9EF03641-91D8-5F34-AA96-850307507501

http://zoobank.org/071280B5-EB72-4445-9605-EAC6699CF789

Figs 28 , 29 , 57D , 67D

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).

Holotype

♂ (IZCAS-Ar 34759, YHCLU0056), : Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, Anogeissus acuminataplantation, 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, AME–AME 0.06, AME–ALE 0.03, PME–PME 0.26, PME–PLE 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

BD53B7FC-8EBB-5729-98F2-4C1306249EF3

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, 48^th 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

F284A349-9331-5877-8B14-7797B83C81C9

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, 48^th 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

5253CF32-E6C2-5C12-9DFD-C8CE21F72D10

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 acuminataplantation, 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

F4DAFE8C-B449-546F-956F-0703732313E1

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

80B4161E-68C0-57DE-BDF9-1E1CA8CD0EF9

• 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

FFF83172-0DA9-51C3-B384-05D657A21DA8

• 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).

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)

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.

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

Figure 58.

Clubiona spp. of the C. filicata group, male palp, ventral view AC. reichliniBC. filicataCC. 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 68.

Clubiona spp. of the C. filicata group, male palp, retrolateral view AC. reichliniBC. filicataCC. banna sp. nov., holotype. Abbreviations: C = conductor; E = embolus; 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 AC. melanostictaBC. zhanggureniCC. circulataDC. 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 AC. grucollarisBC. lalaCC. abnormisDC. yueya. Abbreviations: C = conductor; E = embolus; EB = embolar base; 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 AC. melanostictaBC. zhanggureniCC. circulataDC. 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 AC. grucollarisBC. lalaCC. abnormisDC. yueya. Abbreviations: C = conductor; E = embolus; TA = tegular apophysis; RTA = retrolateral tibial apophysis. Scale bars: 0.1 mm.

Figure 42.

Male palp of Clubiona lalaA 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).

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

Figure 79.

Clubiona spp. of the C. filicata group, epigyne, intact, ventral view AC. circulataBC. reichliniCC. grucollarisDC. lalaEC. suthepicaFC. yueya. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 87.

Clubiona spp. of the C. filicata group, epigyne, cleared, ventral view AC. circulataBC. reichliniCC. grucollarisDC. lalaEC. suthepicaFC. yueya. Abbreviations: A = atrium; AAM = atrial anterior margin; APM = atrial posterior margin; CO = copulatory opening. Scale bars: 0.1 mm.

Figure 95.

Clubiona spp. of the C. filicata group, vulva, cleared, dorsal view AC. circulataBC. reichliniCC. grucollarisDC. lalaEC. suthepicaFC. 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.

Clubiona abnormis

Dankittipakul, 2008

8E4A3C26-C986-5150-8900-FB422361C48F

Figs 30 , 31 , 60C , 70C

Figure 30.

Male palp of Clubiona abnormisA 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).

• 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, 48^th 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.

A12F1FF9-D9DE-5B2D-B77C-EF67ED08CEB3

http://zoobank.org/A821EEFC-C147-4DA6-AE4B-866DD621B2B8

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

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).

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, AME–AME 0.14, AME–ALE 0.07, PME–PME 0.26, PME–PLE 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, AME–AME 0.11, AME–ALE 0.11, PME–PME 0.31, PME–PLE 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

242DBCFF-81F9-5557-99E0-EE9E85B9A403

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

Figure 34.

Male palp of Clubiona circulataA 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).

• 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 banyanplantation, 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).

Figure 40.

Male palp of Clubiona grucollarisA 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 46.

Male palp of Clubiona suthepicaA 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).

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, AME–AME 0.08, AME–ALE 0.04, PME–PME 0.24, PME–PLE 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

D3529FE8-3CCB-5C93-B929-D55897EFECF5

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

Figure 36.

Male palp of Clubiona reichliniA 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).

• 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

A4EB9FD0-D60E-5F68-9CF6-3FB3898BB88F

Figs 38 , 39 , 58B , 68B

Figure 38.

Male palp of Clubiona filicataA 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).

• 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

081BEA46-E6E3-55FC-9B3A-E32E3D4E5E3C

• 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

24630D87-016F-59A2-A4DA-D4BFEFCE6D33

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

47C41D5D-FD27-522E-B6D6-E4FE4DC14F82

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, AME–AME 0.12, AME–ALE 0.10, PME–PME 0.27, PME–PLE 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

04ED1012-F39A-5962-A83E-B43F178036F7

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

Figure 44.

Male palp of Clubiona melanostictaA 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).

• 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 bailloniiplantation (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

1DA253F6-FBCE-5269-8E8E-9770EDFE319B

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, 48^th 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, 48^th 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, AME–AME 0.13, AME–ALE 0.10, PME–PME 0.28, PME–PLE 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

491BF920-D6AC-555F-8242-A20FD162381F

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

6677F0BD-2079-5C22-88F1-2258376890EE

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

16A820C6-8D08-50FC-8D67-A9961EB38603

• 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

1B4E6DE7-D792-59E6-9EA3-9B0A984C21A1

• 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).

Table 6.

Clubionatrivialis 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)

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.

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

F9BE7B31-2B74-5F0F-AC3F-6FC32CB951AD

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

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).

• 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, 48^th 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).

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).

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, AME–AME 0.03, AME–ALE 0.13, PME–PME 0.27, PME–PLE 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

FA335145-C750-57D5-8F0D-1F519AB1D1C5

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.

53DFC2A4-CACB-5C4A-80E1-12DABFB354DE

http://zoobank.org/2CC441EE-1C9B-44EF-8EA7-F575A599F9DD

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, AME–AME 0.04, AME–ALE 0.04, PME–PME 0.20, PME–PLE 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

4BC18759-E76C-5B3A-A023-0F77818D70BA

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, 48^th 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

B3EA9042-59F0-5B5E-ADCF-82608A75FAC9

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 acuminataplantation, 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

F03EB327-03FB-59BE-A58D-358B607D4B6D

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.

8ABA5954-341F-5560-81C5-92AB2096CC7D

http://zoobank.org/442232E1-4DB2-4AA1-86EA-DA3BA9E21858

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

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).

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, AME–AME 0.07, AME–ALE 0.04, PME–PME 0.17, PME–PLE 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, AME–AME 0.08, AME–ALE 0.07, PME–PME 0.23, PME–PLE 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.

41A46993-422D-5745-A54A-F1110A4DF41F

http://zoobank.org/4EB33FD0-CBCB-4642-9B22-78220A539678

Figs 52 , 80C , 88C , 96C

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).

Holotype.

♀ (IZCAS-Ar 34765), China: Yunnan Province: Xishuangbanna: Mengla County: Menglun Town: XTBG, 48^th 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, AME–AME 0.08, AME–ALE 0.11, PME–PME 0.39, PME–PLE 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.

Citation

Zhang J, Yu H, Li S (2021) Taxonomic studies on the sac spider genus Clubiona (Araneae, Clubionidae) from Xishuangbanna Rainforest, China. In: Li S (Ed.) Spiders of Xishuangbanna, China.ZooKeys 1034: 1–163. https://doi.org/10.3897/zookeys.1034.59413