#ZooKeys

ZooKeys 1216: 265-302 (2024)
DOI: 10.3897/zookeys.1216.132561

Research Article

Nine new spider species of Belisana Thorell, 1898 (Araneae,
Pholcidae) from karst caves, with a list of species of the genus
from Guizhou, southwestern China

Bing Wang™, Jinglin Li'®, Shugiang Li?®, Zhiyuan Yao'®

1 College of Life Science, Shenyang Normal University, Shenyang 110034, Liaoning, China
2 Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China
Corresponding authors: Zhiyuan Yao (yaozy@synu.edu.cn); Shugiang Li (lisq@ioz.ac.cn)

OPEN Qaccess

Academic editor: Alireza Zamani
Received: 20 July 2024

Accepted: 3 October 2024
Published: 25 October 2024

ZooBank: https://zoobank.
org/0E173F72-36D3-4B97-BC43-
F096067A3D1E

Citation: Wang B, Li J, Li S, Yao Z
(2024) Nine new spider species of
Belisana Thorell, 1898 (Araneae,
Pholcidae) from karst caves, with a list
of species of the genus from Guizhou,
southwestern China. ZooKeys 1216:
269-302. https://doi.org/10.3897/
zookeys.1216.132561

Copyright: © Bing Wang et al.

This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International - CC BY 4.0).

Abstract

Species of the spider genus Belisana Thorell, 1898 exhibit high diversity in Guizhou,
southwestern China. Previously, only eight species of Belisana were recorded in
Guizhou. In this study, nine new species are described from karst caves: Belisana bijie
Wang, S. Li & Yao, sp. nov., B. /iupanshui Wang, S. Li & Yao, sp. nov., B. majiang Wang,
S. Li & Yao, sp. nov., B. nayong Wang, S. Li & Yao, sp. nov., B. gixingguan Wang, S. Li &
Yao, sp. nov., B. xiuwen Wang, S. Li & Yao, sp. nov., B. yongcong Wang, S. Li & Yao, sp.
nov., B. zhouxi Wang, S. Li & Yao, sp. nov., and B. zunyi Wang, S. Li & Yao, sp. nov. Belis-
ana zhangi Tong & Li, 2007 is reported from Guizhou for the first time. In addition, a list
of all species of Belisana from Guizhou is provided.

Key words: Biodiversity, cellar spiders, daddy-long-legs, invertebrate, morphology,
taxonomy

Introduction

Belisana Thorell, 1898, the second largest genus in the Pholcidae C.L. Koch,
1850, comprises 160 species (WSC 2024). These species inhabit diverse mi-
cro-habitats, e.g., beneath rocks, in caves, on the underside of leaves, among
leaf litter, and amidst foliage in the canopy (Huber 2005; Yao et al. 2015; Zhao
et al. 2023a). They are primarily distributed in southern China, as well as in
the Indo-Malayan and Australasian regions (Huber 2005; Yao and Li 2013; Yao
et al. 2018; Zhu et al. 2020; Zhu and Li 2021). Currently, 46% of the species
(74 spp.) have been documented from southern China (WSC 2024), of which
Yunnan boasts the highest concentration of species, accounting for 42% (31
spp.; Zhang et al. 2024b). Guangxi and Hainan, which have the second and third
highest species diversity of Belisana, respectively, have recorded only 11 and
ten species. Recently, numerous surveys targeting pholcid spiders have been
conducted in China, resulting in the discovery and reporting of a large number
of new species (e.g., Yao et al. 2021; Lu et al. 2022; Zhao et al. 2023b; Yang
et al. 2024a, b). Nevertheless, these efforts have primarily focused on Phol-
cus Walckenaer, 1805, found in epigean environments in northern and central

265

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

China, with relatively few reports on Belisana from hypogean environments in
southern China (10 spp.; Wang et al. 2024; Zhang et al. 2024a, b).

Guizhou, located in the southwest of China, is renowned for its abundant
karst caves. The extreme cave environmental conditions have been regarded
as the primary factors of maintaining species endemism within caves (Lefé-
bure et al. 2006; Yao et al. 2016). Nevertheless, only eight endemic species of
Belisana have been recorded from Guizhou (Chen et al. 2009; Zhang and Peng
2011; Chen et al. 2016; Yang et al. 2023a; Wang et al. 2024). Among these,
seven species are collected from caves. This work aims to report the nine new
species, a new record found in karst caves (Fig. 1), and to provide an updated
list of species of Belisana from Guizhou (Table 1).

Materials and methods

Specimens were examined and measured with a Leica M205 C stereomicro-
scope. Left male palps were photographed. Epigynes were photographed be-
fore dissection. Vulvae were photographed after treating them in a 10% warm
solution of potassium hydroxide (KOH) to dissolve soft tissues. Images were
captured with a Canon EOS 750D wide zoom digital camera (24.2 megapixels)
mounted on the stereomicroscope mentioned above and assembled using Hel-
icon Focus v. 3.10.3 image stacking software (Khmelik et al. 2005). Drawings
were done with Procreate 5.0.2 (Savage Interactive Pty. Ltd.). All measurements
are given in millimeters (mm). Leg measurements are shown as: total length (fe-
mur, patella, tibia, metatarsus, tarsus). Leg segments were measured on their
dorsal sides. The distribution map was generated with ArcGIS v. 10.2 (ESRI
Inc.). The specimens studied are preserved in 75% ethanol and deposited in the
Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing, China.

Table 1. A list of species of Belisana from Guizhou, China.

No. Species Habitat Reference

1 B. bijie sp. nov. karst cave this paper

2 |B. daji Chen, Zhang & Zhu, 2009 karst cave Chen et al. (2009)

3 | B. douging Chen, Zhang & Zhu, 2009 karst cave Chen et al. (2009)

4 |B. galeiformis Zhang & Peng, 2011 / Zhang and Peng (2011)
5 |B. lii Chen, Yu & Guo, 2016 karst cave Chen et al. (2016)

6 _ B. liupanshui sp. nov. karst cave this paper

7 |B. majiang sp. nov. karst cave this paper

8 | B. nayong sp. nov. karst cave this paper

9 |B. qixingguan sp. nov. karst cave this paper

10 |B. wangchengi Wang, Yao & Zhang, 2024 | karst cave Wang et al. (2024)
11. | B. xishui Chen, Zhang & Zhu, 2009 karst cave Chen et al. (2009)
12 |B. xiuwen sp. nov. karst cave this paper

13 |B. yanhe Chen, Zhang & Zhu, 2009 karst cave Chen et al. (2009)
14 |B. yongcong sp. nov. karst cave this paper

15 | B. yuhaoi Yang & Yao, 2023 karst cave Yang et al. (2023a)
16 |B. zhangi Tong & Li, 2007 karst cave this paper; Tong and Li (2007)
17 |B. zhouxi sp. nov. karst cave this paper

18 |B. zunyi sp. nov. karst cave this paper

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561

266

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

q TC
oy ew ve
Guizhou , 27°
o
© O17
7
| Yunnan
150 ki 1 °
Guangxi ear ge \ Be
104° 107° 108° 109°

Figure 1. Distribution records of the Belisana species from Guizhou, China 1 Belisana bijie sp. nov. 2 B. daji Chen, Zhang
& Zhu, 2009 3 B. douging Chen, Zhang & Zhu, 2009 4 B. galeiformis Zhang & Peng, 2011 5 B. lii Chen, Yu & Guo, 2016
6 B. liupanshui sp. nov. 7 B. majiang sp. nov. 8 B. nayong sp. nov. 9 B. gixingguan sp. nov. 10 B. wangchengi Wang, Yao &
Zhang, 2024 11 B. xishui Chen, Zhang & Zhu, 2009 12 B. xiuwen sp. nov. 13 B. yanhe Chen, Zhang & Zhu, 2009 14 B. yong-
cong sp. nov. 15 B. yuhaoi Yang & Yao, 2023 16 B. zhangi Tong & Li, 2007 17 B. zhouxi sp. nov. 18 B. zunyi sp. nov.

Terminology and taxonomic descriptions follow Huber (2005) and Yao et al.
(2015). The following abbreviations are used: aa = anterior arch, ALE = anterior
lateral eye, b = bulb, ba = bulbal apophysis, da = distal apophysis, e = embolus,
ep = epigynal pocket, f = flap, L/d = length/diameter, pa = proximo-lateral apoph-
ysis, PME = posterior median eye, pp = pore plate, pr = procursus.

Taxonomy

Family Pholcidae C.L. Koch, 1850
Subfamily Pholcinae C.L. Koch, 1850

Genus Belisana Thorell, 1898

Type species. Belisana tauricornis Thorell, 1898.

Belisana bijie Wang, S. Li & Yao, sp. nov.
https://Zoobank.org/938091C8-867C-4607-9EB1-9B4BE301F78F
Figs 2, 3, 22A, B, 24A, B

Type material. Holotype: CHINA * 3; Guizhou, Bijie, Qixingguan District, Salaxi
Town, Shuanglongdao Cave; 27°11.493'N, 105°03.850'E; alt. 1920 m; 18 Nov.
2011; Z. Chen & Z. Zha leg.; IZCAS-Ar45175. Paratype: CHINA * 12; same data
as for holotype; IZCAS-Ar45176.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 267

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 2. Belisana bijie sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at ventral
apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow 1
points at prolatero-subdistal sclerite, arrow 2 points at prolatero-distal membranous lamella D retrolateral view, arrow
1 points at sclerotized dorso-subdistal apophysis, arrow 2 points at retrolatero-distal spine, arrow 3 points at retrolate-
ro-subdistal membranous process). Abbreviations: b = bulb, ba = bulbal apophysis, e = embolus, f = flap, pr = procursus.
Scale bars: 0.20 (A, B); 0.05 (C, D).

Diagnosis. The new species resembles B. wangchengi Wang, Yao & Zhang,
2024 (Wang et al. 2024: 2, figs 1A—-D, 2A—H, 3A-D) by having similar male cheli-
cerae (tips of distal apophyses widely separated and pointing inwards; Fig. 3D),
bulbal apophysis (hooked; Fig. 3C), and epigyne (epigynal pockets on lateral
part of epigynal plate; Figs 3A, 24A), but can be distinguished by procursus with
retrolatero-subdistal membranous process (arrow 3 in Figs 2D, 22B vs absent),
by vulval pore plates with nearly angular sclerites (arrow in Figs 3B, 24B vs

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 268

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 3. Belisana bijie sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view B vulva,
dorsal view, arrow points at lateral sclerite C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal view
F lateral view H ventral view). Abbreviations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophysis,
e = embolus, ep = epigynal pocket, pa = proximo-lateral apophysis, pp = pore plate. Scale bars: 0.10 (A-D); 0.40 (E-H).

blunt), and by dorsal shield of prosoma with lateral brown bands (Fig. 3E, G vs
absent, but with median radiating marks).

Description. Male (holotype): Total length 1.95 (2.08 with clypeus), prosoma
0.70 long, 0.82 wide, opisthosoma 1.25 long, 0.90 wide. Leg |: 19.96 (5.13, 0.38,
5.05, 7.52, 1.88), leg Il: 13.25 (3.52, 0.37, 3.32, 4.70, 1.34), leg Ill: 7.81 (2.25,
0.30, 1.90, 2.59, 0.77), leg IV: 9.83 (3.09, 0.30, 2.45, 3.16, 0.83); tibia | L/d: 73.
Eye interdistances and diameters: PME-PME 0.11, PME 0.10, PME-ALE 0.02.
Sternum width/length: 0.63/0.56. Habitus as in Fig. 3E, F. Dorsal shield of pro-
soma yellowish, with brown lateral bands; clypeus brown; sternum yellowish.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 269

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Legs whitish, without darker rings. Opisthosoma yellowish, with black spots.
Thoracic furrow absent. Clypeus unmodified. Chelicerae with pair of proxi-
mo-lateral apophyses (pa in Fig. 3D) and pair of curved distal apophyses (dis-
tance between tips: 0.31; da in Fig. 3D). Palp as in Fig. 2A, B; trochanter with
ventral apophysis (2x longer than wide, arrow 1 in Fig. 2B); femur with retrolat-
ero-proximal protrusion (arrow 2 in Fig. 2B); procursus simple proximally but
complex distally, with prolatero-subdistal sclerite (arrow 1 in Figs 2C, 22A), pro-
latero-distal membranous lamella (arrow 2 in Figs 2C, 22A) bearing proximally
sclerotized part, sclerotized dorso-subdistal apophysis (arrow 1 in Figs 2D, 22B),
retrolatero-distal spine (arrow 2 in Figs 2D, 22B), retrolatero-subdistal membra-
nous process (arrow 3 in Figs 2D, 22B), and retrolateral membranous flap (f in
Figs 2D, 22B); bulb with hooked apophysis (ba in Fig. 3C) and simple embolus
(e in Fig. 3C). Retrolateral trichobothria on tibia | at 9% proximally; legs with
short vertical setae on metatarsi; tarsus | with 17 distinct pseudosegments.

Female (paratype, IZCAS-Ar45176): Similar to male, habitus as in Fig. 3G, H.
Total length 1.96 (2.09 with clypeus), prosoma 0.66 long, 0.83 wide, opistho-
soma 1.30 long, 1.01 wide; tibia |: 3.40; tibia | L/d: 49. Eye interdistances and
diameters: PME-PME 0.10, PME 0.10, PME-ALE 0.02. Sternum width/length:
0.60/0.53. Dorsal shield of prosoma with distinct postero-median marks; cly-
peus yellowish. Epigyne simple and flat, posteriorly curved, with pair of lateral
pockets 0.31 apart (ep in Figs 3A, 24A). Vulva with ridge-shaped anterior arch
bearing pair of nearly angular sclerites (arrow in Figs 3B, 24B), and pair of near-
ly triangular pore plates (4x longer than wide, pp in Figs 3B, 24B).

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guizhou, type locality; Fig. 1).

Etymology. The specific name refers to the type locality; noun in apposition.

Belisana liupanshui Wang, S. Li & Yao, sp. nov.
https://Zzoobank.org/4C5D21CD-86E7-43FA-B974-0235DB601B2B
Figs 4, 5, 22C, D, 24C, D

Type material. Holotype: CHINA + <4; Guizhou, Liupanshui, Pan County, Biyun
Cave; 25°46.527'N, 104°38.278'E; alt. 1468 m; 13 Apr. 2007; J. Liu & Y. Lin
leg.; IZCAS-Ar45181. Paratypes: CHINA - 53; same data as for holotype;
IZCAS-Ar45182-86 * 72; same data as for holotype; IZCAS-Ar45187-93.
Diagnosis. The new species resembles B. jiuxiang Zhang, Li& Yao, 2024 (Zhang
et al. 2024b: 261, figs 4A—D, 5A-H, 18C, D, 20C, D) by having similar male che-
licerae (distal apophyses directed towards frontally, but tips pointing inwards;
Fig. 5D) and epigyne (epigynal pockets on median part of epigynal plate, epigy-
nal plate posteriorly straight; Figs 5A, 24C), but can be distinguished by procur-
sus with prolatero-subdistal and subdistal membranous processes (arrows 1,
3 in Figs 4C, 22C vs absent) and nearly rectangle dorso-subdistal membranous
process (arrow 4 in Figs 4C, 22C vs angular), by bulbal apophysis with angular
subdistal apophysis (arrow in Fig. 5C vs absent), and by vulval pore plates an-
teriorly narrow and posteriorly wide (pp in Figs 5B, 24D vs nearly quadrilateral).
Description. Male (holotype): Total length 1.71 (1.79 with clypeus), prosoma
0.63 long, 0.67 wide, opisthosoma 1.08 long, 0.81 wide. Leg I: — (4.04, 0.29,
4.50, 5.96, —), leg Il: 11.08 (2.80, 0.30, 2.97, 3.92, 1.09), leg III: 7.98 (2.22, 0.25,

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 270

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 4. Belisana liupanshui sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at ven-
tral apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow 1
points at prolatero-subdistal membranous process, arrow 2 points at sclerotized distal apophysis, arrow 3 points at sub-
distal membranous process, arrow 4 points at dorso-subdistal membranous process D retrolateral view). Abbreviations:
b = bulb, ba = bulbal apophysis, e = embolus, f = flap, pr = procursus. Scale bars: 0.10 (A, B); 0.05 (C, D).

1.98, 2.72, 0.81), leg IV: 10.20 (3.00, 0.26, 2.63, 3.45, 0.86); tibia | L/d: 63. Eye
interdistances and diameters: PME-PME 0.09, PME 0.08, PME-ALE 0.02. Ster-
num width/length: 0.51/0.44. Habitus as in Fig. 5E, F. Dorsal shield of prosoma
yellowish, with brownish radiating marks; clypeus brownish; sternum yellowish.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 271

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 5. Belisana liupanshui sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view
B vulva, dorsal view, arrow points at lateral sclerite C bulb, prolateral view, arrow points at subdistal apophysis D chelicer-
ae, frontal view E-H habitus (E, G dorsal view F lateral view H ventral view). Abbreviations: aa = anterior arch, b = bulb, ba
= bulbal apophysis, da = distal apophysis, e = embolus, ep = epigynal pocket, pa = proximo-lateral apophysis, pp = pore
plate. Scale bars: 0.10 (A-D); 0.50 (E-H).

Legs whitish, without darker rings. Opisthosoma yellowish, without spots. Tho-
racic furrow absent. Clypeus unmodified. Chelicerae with pair of proximo-lat-
eral apophyses (pa in Fig. 5D) and pair of curved distal apophyses (distance
between tips: 0.17; da in Fig. 5D). Palp as in Fig. 4A, B; trochanter with ventral

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 272

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

apophysis (as long as wide, arrow 1 in Fig. 4B); femur with retrolatero-proximal
protrusion (arrow 2 in Fig. 4B); procursus simple proximally but complex dis-
tally, with prolatero-subdistal membranous process (arrow 1 in Figs 4C, 22C),
sclerotized distal apophysis (arrow 2 in Figs 4C, 22C), subdistal membranous
process (arrow 3 in Figs 4C, 22C), dorso-subdistal membranous process (arrow
4 in Figs 4C, 22C), and retrolateral membranous flap (f in Figs 4D, 22D); bulb
with hooked apophysis bearing angular subdistal apophysis (arrow in Fig. 5C)
and simple embolus (e in Fig. 5C). Retrolateral trichobothria on tibia | at 4%
proximally; legs with short vertical setae on metatarsi.

Female (paratype, IZCAS-Ar45187): Similar to male, habitus as in Fig. 5G,
H. Total length 2.53 (2.65 with clypeus), prosoma 0.67 long, 0.75 wide, opist-
hosoma 1.86 long, 1.80 wide; tibia |: 4.05; tibia | L/d: 51. Eye interdistances
and diameters: PME-PME 0.11, PME 0.07, PME-ALE 0.02. Sternum width/
length: 0.52/0.46. Epigyne simple and flat, posteriorly straight, with pair of me-
dian pockets 0.20 apart (ep in Figs 5A, 24C). Vulva with ridge-shaped anterior
arch bearing pair of angular lateral sclerites (arrow in Figs 5B, 24D), and pair of
anteriorly narrow and posteriorly wide pore plates (3x longer than wide, pp in
Figs 5B, 24D).

Variation. Tibia | in five male paratypes (IZCAS-Ar45182-86): 4.50, 4.55,
4.62, 4.65, 4.70. Tibia | in the other six female paratypes (IZCAS-Ar45188-93):
3.56=3:85:

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guizhou, type locality; Fig. 1).

Etymology. The specific name refers to the type locality; noun in apposition.

Belisana majiang Wang, S. Li & Yao, sp. nov.
https://Zzoobank.org/75CODB39-2F6A-40D1-93A0-3C85CD956A53
Figs 6, 7, 22E, F, 24E, F

Type material. Holotype: CHINA « 3; Guizhou, Kaili, Majiang County, Xingshan
Town, Gubin Village, Guazhutou Cave; 26°30.257'N, 107°30.943'E; alt. 1056 m;
28 Nov. 2011; Z. Chen & Z. Zha leg.; IZCAS-Ar45194. Paratypes: CHINA * 103;
same data as for holotype; IZCAS-Ar45195 + 29; same data as for holotype;
IZCAS-Ar45196-97.

Diagnosis. The new species resembles B. zhouxi sp. nov. (Figs 18, 19, 23E,
F, 25G, H) by having similar male chelicerae (tips of distal apophyses pointing
downwards; Fig. 7D), bulbal apophysis (hooked; Fig. 7C), and epigyne (epigy-
nal pockets on antero-lateral part of epigynal plate, epigynal plate posterior-
ly curved; Figs 7A, 24E), but can be distinguished by procursus with distinct
ventro-subdistal membranous process and dorso-distal spine (arrows 1, 3 in
Figs 6C, 22E vs absent) and by vulval pore plates long elliptic (3x longer than
wide, pp in Figs 7B, 24F vs 2x); also distinguished from B. tongle Zhang, Chen
& Zhu, 2008 (Zhang et al. 2008: 654, figs 1-5) by procursus with dorso-subdis-
tal membranous process (arrow 4 in Figs 6C, 22E vs absent), by prolatero-dis-
tal membranous lamella of procursus without rectangular sclerite (arrow 2 in
Figs 6C, 22E vs present), by ventro-subdistal membranous process of procur-
sus 3x longer than wide (arrow 1 in Figs 6C, 22E vs 8x), and by dorso-distal
spine of procursus 6x longer than wide (arrow 3 in Figs 6C, 22E vs 12x).

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 273

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 6. Belisana majiang sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at ven-
tral apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow 1
points at ventro-subdistal membranous process, arrow 2 points at prolatero-distal membranous lamella, arrow 3 points
at dorso-distal spine, arrow 4 points at dorso-subdistal membranous process D retrolateral view). Abbreviations: b = bulb,
ba = bulbal apophysis, e = embolus, f = flap, pr = procursus. Scale bars: 0.10 (A, B); 0.02 (C, D).

Description. Male (holotype): Total length 1.35 (1.43 with clypeus), prosoma
0.52 long, 0.59 wide, opisthosoma 0.83 long, 0.64 wide. Leg | missing, leg II:
6.13 (1.78, 0.22, 1.50, 1.90, 0.73), leg Ill: 4.19 (1.28, 0.20, 0.95, 1.33, 0.43), leg
IV: 5.87 (1.72, 0.22, 1.48, 1.80, 0.65). Eye interdistances and diameters: PME-
PME 0.08, PME 0.06, PME-ALE 0.02. Sternum width/length: 0.44/0.38. Habitus

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 274

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 7. Belisana majiang sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view B vulva,
dorsal view C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal view F lateral view H ventral view).
Abbreviations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophysis, e = embolus, ep = epigynal pock-
et, pa = proximo-lateral apophysis, pp = pore plate. Scale bars: 0.05 (A-D); 0.30 (E-H).

as in Fig. 7E, F. Dorsal shield of prosoma yellowish, without marks; clypeus
and sternum yellowish. Legs whitish, without darker rings. Opisthosoma yel-
lowish, without spots. Thoracic furrow absent. Clypeus unmodified. Chelicerae
with pair of proximo-lateral apophyses (pa in Fig. 7D) and pair of curved distal
apophyses (distance between tips: 0.19; da in Fig. 7D). Palp as in Fig. 6A, B;

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 275

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

trochanter with ventral apophysis (1.5x longer than wide, arrow 1 in Fig. 6B);
femur with retrolatero-proximal protrusion (arrow 2 in Fig. 6B); procursus sim-
ple proximally but complex distally, with ventro-subdistal membranous pro-
cess (arrow 1 in Figs 6C, 22E), prolatero-distal membranous lamella (arrow 2
in Figs 6C, 22E) bearing proximally slightly sclerotized part, dorso-distal spine
(arrow 3 in Figs 6C, 22E), dorso-subdistal membranous process (arrow 4 in Figs
6C, 22E), and retrolateral membranous flap (f in Figs 6D, 22F); bulb with hooked
apophysis (ba in Fig. 7C) and simple embolus (e in Fig. 7C).

Female (paratype, IZCAS-Ar45196): Similar to male, habitus as in Fig. 7G, H.
Total length 1.48 (1.58 with clypeus), prosoma 0.48 long, 0.56 wide, opistho-
soma 1.00 long, 0.84 wide. Eye interdistances and diameters: PME-PME 0.08,
PME 0.06, PME-ALE 0.02. Sternum width/length: 0.43/0.39. Epigyne simple
and flat, posteriorly curved, with pair of antero-lateral pockets 0.14 apart (ep
in Figs 7A, 24E). Vulva with ridge-shaped anterior arch (aa in Figs 7B, 24F) and
pair of long elliptic pore plates (3x longer than wide, pp in Figs 7B, 24F).

Variation. Unknown. Legs | missing in male paratype (IZCAS-Ar45195) and
another female paratype (IZCAS-Ar45197).

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guizhou, type locality; Fig. 1).

Etymology. The specific name refers to the type locality; noun in apposition.

Belisana nayong Wang, S. Li & Yao, sp. nov.
https://zoobank.org/2374804C-1CDB-4013-885E-336618ED0D2B
Figs 8, 9, 22G, H, 24G, H

Type material. Holotype: CHINA + <; Guizhou, Bijie, Nayong County, Laosiba
Town, Bailong Cave; 26°50.166'N, 105°31.222'E; alt. 1468 m; 27 Apr. 2007; J.
Liu & Y. Lin leg.; IZCAS-Ar45198. Paratypes: CHINA * 3); same data as for holo-
type; IZCAS-Ar45199-201 * 49; same data as for holotype; IZCAS-Ar45202-05.
Diagnosis. The new species resembles B. yongcong sp. nov. (Figs 14, 15,
23A, B, 25C, D) by having similar bulbal apophysis (hooked; Fig. 9C) and vulva
(anterior arch ridge-shaped, pore plates curved, long elliptic and 8x longer than
wide; Figs 9B, 24H), but can be distinguished by procursus with pointed ven-
tro-subdistal membranous process (arrow 1 in Figs 8C, 22G vs blunt), by male
cheliceral distal apophyses pointing inwards (da in Fig. 9D vs outwards), and by
epigyne with postero-median pockets (ep in Figs 9A, 24G vs lateral).
Description. Male (holotype): Total length 2.33 (2.50 with clypeus), prosoma
0.86 long, 0.89 wide, opisthosoma 1.47 long, 1.15 wide. Leg |: 31.51 (8.33, 0.42,
7.82, 13.08, 1.86), leg II missing, leg III: 19.77 (5.08, 0.39, 5.04, 7.88, 1.38), leg IV
missing; tibia | L/d: 89. Eye interdistances and diameters: PME-PME 0.10, PME
0.10, PME-ALE 0.02. Sternum width/length: 0.68/0.64. Habitus as in Fig. 9E, F.
Dorsal shield of prosoma yellowish, with brown radiating marks; clypeus brown;
sternum yellowish. Legs whitish, without darker rings. Opisthosoma yellowish,
without spots. Thoracic furrow absent. Clypeus unmodified. Chelicerae with
pair of proximo-lateral apophyses (pa in Fig. 9D) and pair of curved distal
apophyses (distance between tips: 0.02; da in Fig. 9D). Palp as in Fig. 8A, B;
trochanter with ventral apophysis (as long as wide, arrow 1 in Fig. 8B); femur
with retrolatero-proximal protrusion (arrow 2 in Fig. 8B); procursus simple

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 276

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 8. Belisana nayong sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at ventral
apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow 1
points at ventro-subdistal membranous process, arrow 2 points at prolatero-distal membranous lamella, arrow 3 points
at dorso-distal spine D retrolateral view). Abbreviations: b = bulb, ba = bulbal apophysis, e = embolus, f = flap, pr = procur-
sus. Scale bars: 0.20 (A, B); 0.05 (C, D).

proximally but complex distally, with ventro-subdistal membranous process
(arrow 1 in Figs 8C, 22G), prolatero-distal membranous lamella (arrow 2 in
Figs 8C, 22G) bearing proximally sclerotized part, dorso-distal spine (arrow 3
in Figs 8C, 22G), and retrolateral membranous flap (f in Figs 8D, 22H); bulb with

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 277

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

a i i an

Figure 9. Belisana nayong sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view B vulva,
dorsal view C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal view F lateral view H ventral view).
Abbreviations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophysis, e = embolus, ep = epigynal pock-
et, pa = proximo-lateral apophysis, pp = pore plate. Scale bars: 0.05 (A-D); 0.40 (E-H).

hooked apophysis (ba in Fig. 9C) and simple embolus (e in Fig. 9C). Retrolateral
trichobothria on tibia | at 4% proximally; legs with short vertical setae on meta-
tarsi; tarsus | with 18 distinct pseudosegments.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 278

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Female (paratype, IZCAS-Ar45202): Similar to male, habitus as in Fig. 9G, H.
Total length 2.47 (2.67 with clypeus), prosoma 0.81 long, 0.88 wide, opistho-
soma 1.66 long, 1.40 wide; tibia |: 4.65; tibia | L/d: 58. Eye interdistances and
diameters: PME-PME 0.11, PME 0.10, PME-ALE 0.02. Sternum width/length:
0.68/0.64. Epigyne simple and flat, posteriorly curved, with pair of postero-me-
dian pockets 0.02 apart (ep in Figs 9A, 24G). Vulva with ridge-shaped anterior
arch (aa in Figs 9B, 24H) and pair of curved, long elliptic pore plates (8x longer
than wide, pp in Figs 9B, 24H).

Variation. Tibia | in three male paratypes (IZCAS-Ar45199-201): 6.15, 6.35,
6.50. Tibia | in the other three female paratypes (IZCAS-Ar45203-05): 4.45,
4.50, 4.74.

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guizhou, type locality; Fig. 1).

Etymology. The specific name refers to the type locality; noun in apposition.

Belisana qixingguan Wang, S. Li & Yao, sp. nov.
https://zoobank.org/36AE89A6-1 3DA-416B-9C06-C86DD3300EC4
Figs 10, 11, 22l, J, 241, J

Type material. Holotype: CHINA * 3; Guizhou, Bijie, Qixingguan District, Chang-
chun Town, Changchun Village, Changchun Cave; 27°13.904'N, 105°10.397'E;
alt. 1580 m; 29 Apr. 2007; J. Liu & Y. Lin leg.; IZCAS-Ar45206. Paratypes: CHINA
- 72; same data as for holotype; IZCAS-Ar45207-13.

Diagnosis. The new species resembles B. tongi Zhang, Li & Yao, 2024 (Zhang
et al. 2024b: 273, figs 12A—D, 13A-H, 18K, L, 21C, D) by having similar dis-
tal part of procursus (distal membranous process nearly half-round; arrow 1
in Figs 10C, 221), bulbal apophysis (hooked; Fig. 11C), and epigyne (epigynal
pockets on postero-lateral part of epigynal plate, epigynal plate posterior-
ly curved; Figs 11A, 241), but can be distinguished by procursus without dis-
tal membranous lamella (Figs 10C, 221 vs present), by male cheliceral distal
apophyses pointing downwards (da in Fig. 11D vs outwards), by vulva without
teeth (Figs 11B, 24J vs present), and by vulval pore plates medially narrow and
posteriorly strongly widened (pp in Figs 11B, 24J vs nearly triangular).

Description. Male (holotype): Total length 1.78 (1.89 with clypeus), prosoma
0.68 long, 0.75 wide, opisthosoma 1.10 long, 0.77 wide. Leg |: 14.43 (3.80, 0.31,
3.72, 5.00, 1.60), leg Il: — (3.00, 0.31, 2.66, 3.75, —), leg III missing, leg IV: 7.17
(2.13, 0.27, 1.66, 2.41, 0.70); tibia | L/d: 50. Eye interdistances and diameters:
PME-PME 0.14, PME 0.08, PME-ALE 0.02. Sternum width/length: 0.55/0.48.
Habitus as in Fig. 11E, F. Dorsal shield of prosoma yellowish, with brown radiat-
ing marks; ocular area with brown median stripe; clypeus brown; sternum yel-
lowish. Legs whitish, without darker rings. Opisthosoma yellowish, with black
spots. Thoracic furrow absent. Clypeus unmodified. Chelicerae with pair of
proximo-lateral apophyses (pa in Fig. 11D) and pair of curved distal apophyses
(distance between tips: 0.29; da in Fig. 11D). Palp as in Fig. 10A, B; trochanter
with ventral apophysis (as long as wide, arrow 1 in Fig. 10B); femur with retro-
latero-proximal protrusion (arrow 2 in Fig. 10B); procursus simple, with distal
membranous process (arrow 1 in Figs 10C, 221) and subdistal spine (arrow 2 in
Figs 10C, 221); bulb with hooked apophysis (ba in Fig. 11C) and simple embolus

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 279

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

pet

Figure 10. Belisana gixingguan sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at
ventral apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow
1 points at distal membranous process, arrow 2 points at subdistal spine D retrolateral view). Abbreviations: b = bulb, ba
= bulbal apophysis, e = embolus, pr = procursus. Scale bars: 0.10 (A, B); 0.05 (C, D).

(e in Fig. 11C). Retrolateral trichobothria on tibia | at 10% proximally; legs with
short vertical setae on metatarsi; tarsus | with 15 distinct pseudosegments.
Female (paratype, IZCAS-Ar45207): Similar to male, habitus as in Fig. 11G,
H. Total length 2.04 (2.15 with clypeus), prosoma 0.78 long, 0.81 wide, opist-
hosoma 1.26 long, 1.00 wide; tibia |: 3.08; tibia | L/d: 39. Eye interdistances

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 280

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 11. Belisana gixingguan sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view B vulva,
dorsal view, arrow points at sclerotized protrusion C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal
view F lateral view H ventral view). Abbreviations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophy-
sis, e = embolus, ep = epigynal pocket, pa = proximo-lateral apophysis, pp = pore plate. Scale bars: 0.10 (A-D); 0.50 (E-H).

and diameters: PME-PME 0.12, PME 0.08, PME-ALE 0.02. Sternum width/
length: 0.55/0.46. Epigyne simple and flat, posteriorly curved, with pair of pos-
tero-lateral pockets 0.28 apart (ep in Figs 11A, 241). Vulva with curved anteri-
or arch (aa in Figs 11B, 24J), pair of medially narrow and posteriorly strongly
widened pore plates (pp in Figs 11B, 24J), and pair of sclerotized protrusions
(arrow in Figs 11B, 24J).

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 281

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Variation. Tibia | in the other six female paratypes (IZCAS-Ar45208-13):
2.89-3.09.

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guizhou, type locality; Fig. 1).

Etymology. The specific name refers to the type locality; noun in apposition.

Belisana xiuwen Wang, S. Li & Yao, sp. nov.
https://zoobank.org/92A64C94-8BCB-421 1-9EDE-4930EEC2467A
Figs 12, 13, 22K, L, 25A, B

Type material. Holotype: CHINA + 3; Guizhou, Guiyang, Xiuwen County, Liutong
Town, Daxing Village, Duobing Cave; 27°06.552'N, 106°29.675'E; alt. 1035 m; 20
Apr. 2007; J. Liu & Y. Lin leg.; IZCAS-Ar45214. Paratypes: CHINA * 39; same data
as for holotype; IZCAS-Ar45215-17.

Diagnosis. The new species resembles B. yongcong sp. nov. (Figs 14, 15, 23A,
B, 25C, D) by having similar male chelicerae (tips of distal apophyses widely
separated and pointing inwards; Fig. 13D), bulbal apophysis (hooked; Fig. 13C),
and epigyne (epigynal pockets on lateral part of epigynal plate, epigynal plate
posteriorly curved; Figs 13A, 25A), but can be distinguished by procursus with
distal spine (arrow 1 in Figs 12C, 22K vs absent) and without ventro-subdistal
membranous lamella, prolatero-distal membranous lamella, and sclerotized
dorso-subdistal apophysis (Figs 12C, 22K vs present), and by vulval pore plates
long elliptic (3x longer than wide, pp in Figs 13B, 25B vs curved and 9x).

Description. Male (holotype): Total length 1.35 (1.48 with clypeus), proso-
ma 0.57 long, 0.59 wide, opisthosoma 0.78 long, 0.60 wide. Leg |: 10.06 (2.48,
0.21, 2.73, 3.36, 1.28), leg Il: 6.94 (2.00, 0.22, 1.74, 2.18, 0.80), leg Ill: 5.08 (1.50,
0.20, 1.14, 1.64, 0.60), leg IV: 6.72 (1.94, 0.20, 1.72, 2.15, 0.71); tibia | L/d: 46.
Eye interdistances and diameters: PME-PME 0.09, PME 0.05, PME-ALE 0.02.
Sternum width/length: 0.45/0.40. Habitus as in Fig. 13E, F. Dorsal shield of pro-
soma yellowish, without marks; clypeus and sternum yellowish. Legs whitish,
without darker rings. Opisthosoma yellowish, without spots. Thoracic furrow
absent. Clypeus unmodified. Eye pigments indistinct. Chelicerae with pair of
proximo-lateral apophyses (pa in Fig. 13D) and pair of curved distal apophyses
(distance between tips: 0.30; da in Fig. 13D). Palp as in Fig. 12A, B; trochanter
with ventral apophysis (as long as wide, arrow 1 in Fig. 12B); femur with retro-
latero-proximal protrusion (arrow 2 in Fig. 12B); procursus simple, with distal
spine (arrow 1 in Figs 12C, 22K) and retrolateral membranous flap (f in Figs 12D,
22L); bulb with hooked apophysis (ba in Fig. 13C) and simple embolus (e in
Fig. 13C). Retrolateral trichobothria on tibia | at 6% proximally; legs with short
vertical setae on metatarsi; tarsus | with 15 distinct pseudosegments.

Female (paratype, IZCAS-Ar45215): Similar to male, habitus as in Fig. 13G,
H. Total length 1.75 (1.85 with clypeus), prosoma 0.56 long, 0.59 wide, opist-
hosoma 1.19 long, 0.80 wide; tibia I: 2.19; tibia | L/d: 44. Eye interdistances and
diameters: PME-PME 0.09, PME 0.05, PME-ALE 0.02. Sternum width/length:
0.44/0.38. Epigyne simple and flat, posteriorly curved, with pair of lateral pock-
ets 0.33 apart (ep in Figs 13A, 25A). Vulva with ridge-shaped anterior arch (aa
in Figs 13B, 25B) and pair of long elliptic pore plates (3x longer than wide, pp
in Figs 13B, 25B).

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 282

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 12. Belisana xiuwen sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at ven-
tral apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow
1 points at distal spine D retrolateral view). Abbreviations: f = flap, pr = procursus. Scale bars: 0.10 (A, B); 0.02 (C, D).

Variation. Tibia | in the other two female paratypes (IZCAS-Ar45216-17):
ZARA US:

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guizhou, type locality; Fig. 1).

Etymology. The specific name refers to the type locality; noun in apposition.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 993

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 13. Belisana xiuwen sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view B vulva,
dorsal view C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal view F lateral view H ventral view).
Abbreviations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophysis, e = embolus, ep = epigynal pock-
et, pa = proximo-lateral apophysis, pp = pore plate. Scale bars: 0.05 (A-D); 0.50 (E-H).

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 284

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Belisana yongcong Wang, S. Li & Yao, sp. nov.
https://zoobank.org/1E1B4058-113E-48F6-9734-39D486FFOFB9
Figs 14, 15, 23A, B, 25C, D

Type material. Holotype: CHINA + 3; Guizhou, Qiandongnan, Liping County,
Yongcong Town, Guantuan Village, Guantuan Cave; 25°59.028'N, 109°06.979'E;
alt. 544 m; 23 May 2007; J. Liu & Y. Lin leg.; IZCAS-Ar45218. Paratypes: CHINA
* 23; same data as for holotype; IZCAS-Ar45219-20 + 59; same data as for
holotype; IZCAS-Ar45221-25.

Diagnosis. The new species resembles B. nayong sp. nov. (Figs 8, 9, 22G, H,
24G, H) by having similar bulbal apophysis (hooked; Fig. 15C) and vulva (anteri-
or arch ridge-shaped, pore plates curved, long elliptic and 8x longer than wide;
Figs 15B, 25D), but can be distinguished by procursus with blunt ventro-sub-
distal membranous process (arrow 1 in Figs 14C, 23A vs pointed), by male
cheliceral distal apophyses pointing outwards (da in Fig. 15D vs inwards), and
by epigyne with lateral pockets (ep in Figs 15A, 25C vs postero-median).

Description. Male (holotype): Total length 1.73 (1.82 with clypeus), prosoma
0.83 long, 0.67 wide, opisthosoma 0.90 long, 0.66 wide. Leg |: 14.76 (3.60, 0.34,
4.05, 5.58, 1.56), leg Il: 12.45 (3.36, 0.32, 3.08, 4.60, 1.09), leg Ill: 8.21 (2.38,
0.26, 1.98, 2.84, 0.75), leg IV: 10.64 (3.20, 0.26, 2.66, 3.72, 0.80); tibia | L/d: 46.
Eye interdistances and diameters: PME-PME 0.06, PME 0.09, PME-ALE 0.02.
Sternum width/length: 0.54/0.49. Habitus as in Fig. 15E, F. Dorsal shield of pro-
soma yellowish, with brownish radiating marks; clypeus and sternum yellowish.
Legs whitish, without darker rings. Opisthosoma yellowish, without spots. Tho-
racic furrow absent. Clypeus unmodified. Chelicerae with pair of proximo-lat-
eral apophyses (pa in Fig. 15D) and pair of curved distal apophyses (distance
between tips: 0.32; da in Fig. 15D). Palp as in Fig. 14A, B; trochanter with ventral
apophysis (as long as wide, arrow 1 in Fig. 14B); femur with retrolatero-prox-
imal protrusion (arrow 2 in Fig. 14B); procursus simple proximally but com-
plex distally, with ventro-subdistal membranous lamella (arrow 1 in Figs 14C,
23A) bearing proximally sclerotized part, prolatero-distal membranous lamel-
la (arrow 2 in Figs 14C, 23A) bearing proximally sclerotized part, sclerotized
dorso-subdistal apophysis (arrow 3 in Figs 14C, 23A), and retrolateral mem-
branous flap (f in Figs 14D, 23B); bulb with hooked apophysis (ba in Fig. 15C)
and simple embolus (e in Fig. 15C). Retrolateral trichobothria on tibia | at 10%
proximally; legs with short vertical setae on metatarsi; tarsus | with 20 distinct
pseudosegments.

Female (paratype, IZCAS-Ar45221): Similar to male, habitus as in Fig. 15G,
H. Total length 2.17 (2.29 with clypeus), prosoma 0.87 long, 0.76 wide, opist-
hosoma 1.30 long, 1.08 wide; tibia I: 3.65; tibia | L/d: 47. Eye interdistances and
diameters: PME-PME 0.09, PME 0.08, PME-ALE 0.02. Sternum width/length:
0.59/0.45. Epigyne simple and flat, posteriorly curved, with pair of lateral pock-
ets 0.35 apart (ep in Figs 15A, 25C). Vulva with ridge-shaped anterior arch (aa in
Figs 15B, 25D) and pair of curved, long elliptic pore plates (9x longer than wide,
pp in Figs 15B, 25D).

Variation. Tibia | in one male paratype (IZCAS-Ar45219): 5.32 (leg | missing in
IZCAS-Ar45220). Tibia lin the other four female paratypes (IZCAS-Ar45222-25):
3.762.3:855369654) 16:

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 285

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 14. Belisana yongcong sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at
ventral apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow
1 points at ventro-subdistal membranous lamella, arrow 2 points at prolatero-distal membranous lamella, arrow 3 points
at sclerotized dorso-subdistal apophysis D retrolateral view). Abbreviations: b = bulb, ba = bulbal apophysis, e = embolus,
f = flap, pr = procursus. Scale bars: 0.10 (A, B); 0.02 (C, D).

Habitat. The species was found in the dark zone inside the cave.
Distribution. China (Guizhou, type locality; Fig. 1).
Etymology. The specific name refers to the type locality; noun in apposition.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 286

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 15. Belisana yongcong sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view
B vulva, dorsal view C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal view F lateral view H ventral
view). Abbreviations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophysis, e = embolus, ep = epigynal
pocket, pa = proximo-lateral apophysis, pp = pore plate. Scale bars: 0.10 (A—D); 0.40 (E-H).

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 287

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Belisana zhangi Tong & Li, 2007
FIgs<16; 17,-23C, De 25ExF

Belisana zhangi Tong and Li 2007: 505, figs 1-6.

Material examined. CHINA: 1; Guizhou, Anshun, Puding County, Huachu Town,
Jingian Cave; 26°14.835'N, 105°37.521'E; 3 May 2005; collector unknown; IZ-
CAS-Ar45177 * 39; same data as for preceding; IZCAS-Ar45178-80.

Diagnosis. The species resembles B. yuhaoi Yang & Yao, 2023 (Yang et al.
2023a: 178, figs 2A, B, 3A—D, 4A-H) by having similar bulbal apophysis (hooked;
Fig. 17C) and epigyne (epigynal pockets on posterior part of epigynal plate,
epigynal plate posteriorly curved; Figs 17A, 25E), but can be distinguished by
procursus with nearly angular ventro-subdistal membranous lamella (arrow 1
in Figs 16C, 23C vs nearly round) and curved retrolateral membranous flap (f in
Figs 16D, 23D vs angular), by male cheliceral distal apophyses pointing inwards
(da in Fig. 17D vs downwards), and by vulval pore plates long elliptic (pp in
Figs 17B, 25F vs nearly triangular).

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guangxi, type locality; Guizhou, Fig. 1).

Belisana zhouxi Wang, S. Li & Yao, sp. nov.
https://zoobank.org/D2E3FODB-7736-46DA-ADC9-9F1FC1CF3ACD
Figs 18, 19, 23E, F, 25G, H

Type material. Holotype: CHINA > 3; Guizhou, Kaili, Zhouxi Town, Hebian
Cave; 26°29.280'N, 107°55.442'E; alt. 665 m; 25 May 2007; J. Liu & Y. Lin
leg.; IZCAS-Ar45226. Paratypes: CHINA * 29°; same data as for holotype; IZ-
CAS-Ar45227-28.

Diagnosis. The new species resembles B. majiang sp. nov. (Figs 6, 7, 22E,
F, 24E, F) by having similar male chelicerae (tips of distal apophyses pointing
downwards; Fig. 19D), bulbal apophysis (hooked; Fig. 19C), and epigyne (epigy-
nal pockets on antero-lateral part of epigynal plate, epigynal plate posteriorly
curved; Figs 19A, 25G), but can be distinguished by procursus without ven-
tro-subdistal membranous process and dorso-distal spine (Figs 18C, 23E vs
present) and by vulval pore plates elliptic (2x longer than wide, pp in Figs 19B,
25H vs 3x).

Description. Male (holotype): Total length 1.17 (1.27 with clypeus), prosoma
0.46 long, 0.50 wide, opisthosoma 0.71 long, 0.61 wide. Leg I: 6.72 (1.84, 0.21,
1.66, 2.13, 0.88), leg Il: 4.56 (1.36, 0.19, 1.06, 1.36, 0.59), leg III: 3.29 (0.96, 0.19,
0.66, 1.01, 0.47), leg IV: 4.69 (1.44, 0.18, 1.16, 1.42, 0.49); tibia | L/d: 27. Eye
interdistances and diameters: PME-PME 0.08, PME 0.06, PME-ALE 0.02. Ster-
num width/length: 0.41/0.33. Habitus as in Fig. 19E, F. Dorsal shield of prosoma
yellowish, without marks; clypeus and sternum yellowish. Legs whitish, without
darker rings. Opisthosoma yellowish, without spots. Thoracic furrow absent.
Clypeus unmodified. Chelicerae with pair of proximo-lateral apophyses (pa in
Fig. 19D) and pair of curved distal apophyses (distance between tips: 0.19; da
in Fig. 19D). Palp as in Fig. 18A, B; trochanter with ventral apophysis (as long
as wide, arrow 1 in Fig. 18B); femur with retrolatero-proximal protrusion (arrow

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 288

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

D

Figure 16. Belisana zhangi Tong & Li, 2007, male A, B palp (A prolateral view B retrolateral view, arrow 1 points at ventral
apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow 1
points at ventro-subdistal membranous lamella, arrow 2 points at prolatero-distal membranous lamella D retrolateral
view, arrow 1 points at sclerotized dorso-subdistal apophysis, arrow 2 points at retrolatero-distal spine). Abbreviations:
b = bulb, ba = bulbal apophysis, e = embolus, f = flap, pr = procursus. Scale bars: 0.10 (A, B); 0.02 (C, D).

2 in Fig. 18B); procursus simple proximally but complex distally, with prolat-
ero-distal membranous lamella (arrow 1 in Figs 18C, 23E) bearing proximal-
ly slightly sclerotized part, dorso-subdistal membranous process (arrow 2 in
Figs 18C, 23E), and retrolateral membranous flap (f in Figs 18D, 23F); bulb with

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 289

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 17. Belisana zhangi Tong & Li, 2007, male (C-F) and female (A, B, G, H) A epigyne, ventral view B vulva, dorsal
view C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal view F lateral view H ventral view). Abbre-
viations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophysis, e = embolus, ep = epigynal pocket, pa
= proximo-lateral apophysis, pp = pore plate. Scale bars: 0.10 (A—D); 0.40 (E-H).

hooked apophysis (ba in Fig. 19C) and simple embolus (e in Fig. 19C). Retro-
lateral trichobothria on tibia | at 7% proximally; legs with short vertical setae on
metatarsi; tarsus | with 14 distinct pseudosegments.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 290

Bing Wang et al.: Ten new species of Belisana spiders, with a list from Guizhou, China

Figure 18. Belisana zhouxi sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at ventral
apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow 1 points
at prolatero-distal membranous lamella, arrow 2 points at dorso-subdistal membranous process D retrolateral view).
Abbreviations: b = bulb, ba = bulbal apophysis, e = embolus, f = flap, pr = procursus. Scale bars: 0.05 (A, B); 0.02 (C, D).

Female (paratype, IZCAS-Ar45227): Similar to male, habitus as in Fig. 19G,
H. Total length 1.21 (1.31 with clypeus), prosoma 0.49 long, 0.57 wide, opist-
hosoma 0.72 long, 0.63 wide; tibia |: 1.22; tibia | L/d: 22. Eye interdistances
and diameters: PME-PME 0.07, PME 0.06, PME-ALE 0.01. Sternum width/

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 291

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

es 1 ANN OO 2G. |
Figure 19. Belisana zhouxi sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view B vulva,
dorsal view C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal view F lateral view H ventral view).
Abbreviations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophysis, e = embolus, ep = epigynal pock-
et, pa = proximo-lateral apophysis, pp = pore plate. Scale bars: 0.05 (A-D); 0.20 (E-H).

length: 0.43/0.32. Epigyne simple and flat, posteriorly curved, with pair of an-
tero-lateral pockets 0.16 apart (ep in Figs 19A, 25G). Vulva with ridge-shaped
anterior arch (aa in Figs 19B, 25H) and pair of elliptic pore plates (2x longer
than wide, pp in Figs 19B, 25H).

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 299

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Variation. Unknown. Leg | missing in another female paratype
(IZCAS-Ar45228).

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guizhou, type locality; Fig. 1).

Etymology. The specific name refers to the type locality; noun in apposition.

Belisana zunyi Wang, S. Li & Yao, sp. nov.
https://zoobank.org/7A299DDB-C1 1 D-4D42-8354-65D16BC50C90
Figs720,2:1) 236-HP25l <3

Type material. Holotype: CHINA * 3; Guizhou, Zunyi, Tongzi County, Leishan-
guan Town, Yangliuping Village; 28°08.692'N, 106°47.378'E; alt. 1008 m; 11
May 2007; J. Liu & Y. Lin leg.; IZCAS-Ar45229. Paratypes: CHINA « 4); same
data as for holotype; IZCAS-Ar45230-33 + 139; same data as for holotype;
IZCAS-Ar45234-46.

Diagnosis. The new species resembles B. zhangi Tong & Li, 2007 (Tong and Li
2007: 505, figs 1-6) by having similar bulbal apophysis (hooked; Fig. 21C) and
vulva (anterior arch ridge-shaped, pore plates long elliptic and 6x longer than
wide; Figs 21B, 25J), but can be distinguished by procursus with dorso-subdis-
tal membranous process (arrow 3 in Figs 20C, 23G vs sclerotized apophysis)
and without ventro-subdistal membranous lamella (Figs 20C, 23G vs present),
by tips of male cheliceral distal apophyses close to each other (da in Fig. 21D
vs widely separated), and by epigyne with median pockets (ep in Figs 21A, 251
vs postero-median).

Description. Male (holotype): Total length 1.62 (1.72 with clypeus), proso-
ma 0.60 long, 0.66 wide, opisthosoma 1.02 long, 0.78 wide. Leg |: 13.53 (3.60,
0.29, 3.50, 4.60, 1.54), leg Il: 8.40 (2.25, 0.30, 2.08, 2.81, 0.96), leg Ill: 5.69 (1.64,
0.25, 1.34, 1.86, 0.60), leg IV: 7.72 (2.12, 0.24, 2.00, 2.56, 0.80); tibia | L/d: 43.
Eye interdistances and diameters: PME-PME 0.09, PME 0.08, PME-ALE 0.02.
Sternum width/length: 0.53/0.47. Habitus as in Fig. 21E, F. Dorsal shield of pro-
soma yellowish, without marks; clypeus and sternum yellowish. Legs whitish,
without darker rings. Opisthosoma yellowish, without spots. Thoracic furrow
absent. Clypeus unmodified. Chelicerae with pair of proximo-lateral apophyses
(pa in Fig. 21D) and pair of curved distal apophyses (distance between tips:
0.04; da in Fig. 21D). Palp as in Fig. 20A, B; trochanter with ventral apophysis
(as long as wide, arrow 1 in Fig. 20B); femur with retrolatero-proximal protru-
sion (arrow 2 in Fig. 20B); procursus simple proximally but complex distally,
with prolatero-distal membranous lamella (arrow 1 in Figs 20C, 23G) bearing
proximally sclerotized part, dorso-distal spine (arrow 2 in Figs 20C, 23G), dor-
so-subdistal membranous process (arrow 3 in Figs 20C, 23G), and retrolater-
al membranous flap (f in Figs 20D, 23H); bulb with hooked apophysis (ba in
Fig. 21C) and simple embolus (e in Fig. 21C). Retrolateral trichobothria on tibia
| at 10% proximally; legs with short vertical setae on metatarsi; tarsus | with 15
distinct pseudosegments.

Female (paratype, IZCAS-Ar45234): Similar to male, habitus as in Fig. 21G,
H. Total length 1.57 (1.66 with clypeus), prosoma 0.60 long, 0.68 wide, opist-
hosoma 0.97 long, 0.78 wide; tibia I: 2.33; tibia | L/d: 35. Eye interdistances and
diameters: PME-PME 0.08, PME 0.06, PME-ALE 0.02. Sternum width/length:

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 293

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 20. Belisana zunyi sp. nov., holotype male A, B palp (A prolateral view B retrolateral view, arrow 1 points at ventral
apophysis, arrow 2 points at retrolatero-proximal protrusion) C, D distal part of procursus (C prolateral view, arrow 1
points at prolatero-distal membranous lamella, arrow 2 points at dorso-distal spine, arrow 3 points at dorso-subdistal
membranous process D retrolateral view). Abbreviations: f = flap, pr = procursus. Scale bars: 0.10 (A, B); 0.03 (C, D).

0.50/0.46. Epigyne simple and flat, posteriorly straight, with pair of median
pockets 0.06 apart (ep in Figs 21A, 251). Vulva with ridge-shaped anterior arch
(aa in Figs 21B, 25J) and pair of long elliptic pore plates (6x longer than wide,
pp in Figs 21B, 25J).

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 294

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 21. Belisana zunyi sp. nov., holotype male (C-F) and paratype female (A, B, G, H) A epigyne, ventral view B vulva,
dorsal view C bulb, prolateral view D chelicerae, frontal view E-H habitus (E, G dorsal view F lateral view H ventral view).
Abbreviations: aa = anterior arch, b = bulb, ba = bulbal apophysis, da = distal apophysis, e = embolus, ep = epigynal pock-
et, pa = proximo-lateral apophysis, pp = pore plate. Scale bars: 0.10 (A-D); 0.30 (E-H).

Variation. Tibia | in four male paratypes (IZCAS-Ar45230—45233): 3.12, 3.20,
3.28, 3.70. Tibia lin the other 12 female paratypes (IZCAS-Ar45235-46): 2.31—2.50.

Habitat. The species was found in the dark zone inside the cave.

Distribution. China (Guizhou, type locality; Fig. 1).

Etymology. The specific name refers to the type locality; noun in apposition.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 295

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 22. Procursus in prolateral and retrolateral views (The arrows point at the same structures as those shown in the
photos of each species) A, B Belisana bijie sp. nov. C, D B. liupanshui sp. nov. E, F B. majiang sp. nov. G, H B. nayong sp.
nov. I, J B. gixingguan sp. nov. K, L B. xiuwen sp. nov. Abbreviation: f = flap. Scale bars: 0.10.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 296

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 23. Procursus in prolateral and retrolateral views (The arrows point at the same structures as those shown in the
photos of each species) A, B Belisana yongcong sp. nov. C, D B. zhangi Tong & Li, 2007 E, F B. zhouxi sp. nov. G, H B. zunyi
sp. nov. Abbreviation: f = flap. Scale bars: 0.10.

Discussion

Belisana is highly diverse in southern China. Including the nine species de-
scribed in this paper, there are now 83 species of Belisana in southern China,
representing 49% of the global total of the genus. Thailand, Indonesia, and Viet-
nam rank second, third, and fourth, respectively, in species diversity of Belisana;
however, these countries have recorded only 19, 18, and 17 species, respec-
tively. Other countries, such as Laos (8 spp.), Malaysia (8 spp.), and Sri Lanka
(6 spp.), have recorded fewer than ten species (WSC 2024). This high level of
activity in China contrasts with the sporadic coverage of Southeast Asia, where
most research has been conducted by foreign arachnologists, and several
countries lack native expertise in this field (Yao and Li 2021; Zhang et al. 2022;
Yang et al. 2023b). Given that Southeast Asia encompasses the Indo-Burma
and Sundaland biodiversity hotspots, we anticipate that further exploration will
reveal additional, as-yet-undiscovered species diversity of Belisana.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 297

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 24. Female genitalia in ventral and dorsal views A, B Belisana bijie sp. nov., arrow points at lateral sclerite C, D B. liupans-
hui sp. nov., arrow points at lateral sclerite E, F B. majiang sp. nov. G, H B. nayong sp. nov I, J B. gixingguan sp. nov., arrow points
at sclerotized protrusion. Abbreviations: aa = anterior arch, ep = epigynal pocket, pp = pore plate. Scale bars: 0.10.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 298

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Figure 25. Female genitalia in ventral and dorsal views A, B Belisana xiuwen sp. nov. C, DB. yongcong sp. nov. E, F B. zhan-
gi Tong & Li, 2007 G, H B. zhouxi sp. nov. I, J B. zunyi sp. nov. Abbreviations: aa = anterior arch, ep = epigynal pocket, pp
= pore plate. Scale bars: 0.10.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 299

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Acknowledgements

The manuscript benefited greatly from comments by Alireza Zamani, Yanfeng
Tong, and Bernhard Huber.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported by the National Natural Science Foundation of China (NSFC-
32170461, 31872193).

Author contributions

ZY and SL designed the study. BW, SL, and ZY performed morphological species identi-
fication. BW and JL finished the descriptions and took the photos and drawings. ZY, BW,
and SL drafted and revised the manuscript.

Author ORCIDs

Bing Wang © https://orcid.org/0009-0009-7047-4395
Jinglin Li © https://orcid.org/0009-0008-3932-309X
Shugiang Li © https://orcid.org/0000-0002-3290-5416
Zhiyuan Yao ® https://orcid.org/0000-0002-1631-0949

Data availability

All of the data that support the findings of this study are available in the main text.

References

Chen H, Zhang F, Zhu M (2009) Four new troglophilous species of the genus Belisana
Thorell, 1898 (Araneae, Pholcidae) from Guizhou Province, China. Zootaxa 2092: 58-
68. https://doi.org/10.11646/zootaxa.2092.1.5

Chen H, Yu Z, Guo X (2016) One new spider species of genus Belisana (Araneae: Pholci-
dae) from Guizhou, China. Sichuan Journal of Zoology 35(2): 227-230.

Huber BA (2005) High species diversity, male-female coevolution, and metaphyly in
Southeast Asian pholcid spiders: The case of Belisana Thorell 1898 (Araneae, Pholci-
dae). Zoologica 155: 1-126.

Khmelik VV, Kozub D, Glazunov A (2005) Helicon Focus 3.10.3. https://www.heliconsoft.
com/heliconsoft-products/helicon-focus/ [Accessed 6 July 2024]

Lefébure T, Douady CJ, Gouy M, Trontelj P Briolay J, Gibert J (2006) Phylogeography of
a subterranean amphipod reveals cryptic diversity and dynamic evolution in extreme
environments. Molecular Ecology 15: 1797-1806. https://doi.org/10.1111/j.1365-
294X.2006.02888.x

Lu Y, Chu C, Zhang X, Li S, Yao Z (2022) Europe vs China: Pholcus (Araneae, Pholcidae)
from Yanshan-Taihang Mountains confirms uneven distribution of spiders in Eurasia.

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 300

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

Zoological Research 43(4): 532-534 [& Suppl. 1-78]. https://doi.org/10.24272/j.
issn.2095-8137.2022.103

Tong Y, Li S (2007) A new six-eyed pholcid spider (Araneae, Pholcidae) from Karst Tian-
keng of Leye County, Guangxi, China. Acta Zootaxonomica Sinica 32: 505-507.

Wang B, Yao Z, Zhang X (2024) A new spider species of Belisana Thorell, 1898 (Araneae,
Pholcidae) from Guizhou Province, south-western China. Biodiversity Data Journal
12: e125111 [1-7]. https://doi.org/10.3897/BDJ.12.e125111

WSC (2024) World Spider Catalog, Version 25.5. Natural History Museum Bern.
https://wsc.nmbe.ch [Accessed 19 September 2024]

Yang L, Zhao F, He Q, Yao Z (2023a) A survey of pholcid spiders (Araneae, Pholci-
dae) from Guiyang, Guizhou Province, China. ZooKeys 1186: 175-184. https://doi.
org/10.3897/zookeys.1186.105736

Yang R, Yan M, Zhang L, Liu H, Koh JKH, He Q, Yao Z (2023b) New taxa of spiders (Ara-
neae) from the world in 2022. Biodiversity Science 31(10): 23175 [1-7]. https://doi.
org/10.17520/biods.2023175

Yang L, Fu C, Zhang Y, He Q, Yao Z (2024a) A survey of Pholcus spiders (Araneae, Phol-
cidae) from the Qinling Mountains of central China, with descriptions of seven new
species. Zoosystematics and Evolution 100(1): 199-221. https://doi.org/10.3897/
zse.100.116759

Yang L, He Q, Yao Z (2024b) Taxonomic study of four closely-related species of the Phol-
cus yichengicus species group (Araneae, Pholcidae) from China’s Qinling Mountains:
An integrated morphological and molecular approach. Zoosystematics and Evolution
100(1): 279-289. https://doi.org/10.3897/zse.100.115633

Yao Z, Li S (2013) New and little known pholcid spiders (Araneae: Pholcidae) from Laos.
Zootaxa 3709(1): 1-51. https://doi.org/10.11646/zootaxa.3709.1.1

Yao Z, Li S (2021) Annual report of Chinese spider taxonomy in 2020. Biodiversity Sci-
ence 29(8): 1058-1063. https://doi.org/10.17520/biods.2021140

Yao Z, Pham DS, Li S (2015) Pholcid spiders (Araneae: Pholcidae) from northern Viet-
nam, with descriptions of nineteen new species. Zootaxa 3909(1): 1-82. https://doi.
org/10.11646/zootaxa.3909.1.1

Yao Z, Dong T, Zheng G, Fu J, Li S (2016) High endemism at cave entrances: a case
study of spiders of the genus Uthina. Scientific Reports 6: 35757 [1-9 & Suppl. I-LIl].
https://doi.org/10.1038/srep35757

Yao Z, Zhu K, Du Z, Li S (2018) The Belisana spiders (Araneae: Pholcidae) from Xish-
uangbanna Tropical Botanical Garden, Yunnan, China. Zootaxa 4425(2): 243-262.
https://doi.org/10.11646/zootaxa.4425.2.3

Yao Z, Wang X, Li S (2021) Tip of the iceberg: species diversity of Pholcus spiders
(Araneae, Pholcidae) in the Changbai Mountains, northeast China. Zoological
Research 42(3): 267-271 [& Suppl. 1-60]. https://doi.org/10.24272/j.issn.2095-
8137.2021.037

Zhang F, Peng Y (2011) Eleven new species of the genus Belisana Thorell (Araneae:
Pholcidae) from south China. Zootaxa 2989: 51-68. https://doi.org/10.11646/zoo-
taxa.2989.1.2

Zhang Y, Chen H, Zhu M (2008) A new troglophilous Belisana spider from Guangxi, China
(Araneae, Pholcidae). Acta Zootaxonomica Sinica 33: 654-656.

Zhang L, Lu Y, Chu C, He Q, Yao Z (2022) New spider taxa of the world in 2021. Biodiver-
sity Science 30(8): 22163 [1-8]. https://doi.org/10.17520/biods.2022163

Zhang L, Wang Y, Li S, Zhang X, Yao Z (2024a) Three new spider species of Belisana
Thorell, 1898 (Araneae, Pholcidae) from karst caves, with a list of Belisana species

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 301

Bing Wang et al.: Nine new species of Belisana spiders, with a list from Guizhou, China

from Guangxi, China. ZooKeys 1209: 315-330. https://doi.org/10.3897/zook-
eys.1209.127951

Zhang L, Wu Z, Li S, Yao Z (2024b) Eight new spider species of Belisana Thorell, 1898
(Araneae, Pholcidae), with an updated overview of Belisana species from Yunnan,
China. ZooKeys 1202: 255-286. https://doi.org/10.3897/zookeys.1202.121633

Zhao F, Yang L, Li S, Zheng G, Yao Z (2023a) A further study on the Belisana spiders (Ara-
neae: Pholcidae) from Xishuangbanna, Yunnan, China. Zootaxa 5351(5): 543-558.
https://doi.org/10.11646/zootaxa.5351.5.3

Zhao F, Yang L, Zou Q, Ali A, LiS, Yao Z (2023b) Diversity of Pholcus spiders (Araneae:
Pholcidae) in China’s Liliang Mountains: an integrated morphological and molecular
approach. Insects 14(4): 364 [1-34]. https://doi.org/10.3390/insects14040364

Zhu W, Li S (2021) Six new species of the spider genus Belisana (Araneae: Pholcidae)
from Southeast Asia. Zootaxa 4963(1): 115-137. https://doi.org/10.11646/zoot-
axa.4963.1.5

Zhu W, Yao Z, Zheng G, Li S (2020) Six new species of the spider genus Belisana
Thorell, 1898 (Araneae: Pholcidae) from southern China. Zootaxa 4810(1): 175-197.
https://doi.org/10.11646/zootaxa.4810.1.12

ZooKeys 1216: 265-302 (2024), DOI: 10.3897/zookeys.1216.132561 302