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Cave-dwelling pseudoscorpions of China 61
Cave-dwelling pseudoscorpions of China with
descriptions of four new hypogean species of Parobisium
(Pseudoscorpiones, Neobisiidae)
from Guizhou Province
Zegang Feng1, J. Judson Wynne2, Feng Zhang1
1 e Key Laboratory of Zoological Systematics and Application, College of Life Sciences, Hebei University,
Baoding, Hebei 071002, China 2 Department of Biological Sciences, Colorado Plateau Museum of Arthropod
Biodiversity and Merriam-Powell Center for Environmental Research, Northern Arizona University, Flagsta,
Arizona 86011, USA
Corresponding author: Feng Zhang (dudu06042001@163.com)
Academic editor: O. Moldovan|Received 23 December 2019|Accepted 22 February 2020|Published 26 March2020
http://zoobank.org/3FD733A3-F576-42AC-9ACA-7CAF915DD82C
Citation: Feng Z, Wynne JJ, Zhang F (2020) Cave-dwelling pseudoscorpions of China with descriptions of four new
hypogean species of Parobisium (Pseudoscorpiones, Neobisiidae) from Guizhou Province. Subterranean Biology 34: 61–98.
https://doi.org/10.3897/subtbiol.34.49586
Abstract
We summarize and discuss the 29 known cave-dwelling pseudoscorpion species from China. Four new
troglomorphic pseudoscorpion species, Parobisium motianense sp. nov., P. qiangzhuang sp. nov., P. san-
louense sp. nov., and P. tiani sp. nov., belonging to the family Neobisiidae, are described based on speci-
mens collected in karst caves in Guizhou, China. Detailed diagnosis, descriptions, and illustrations are
presented. We also provide recommendations for management of caves where they occur, as well as the
cave arthropod communities and the habitats that support them.
Keywords
cavernicoles, cave conservation, taxonomy, troglobionts
Subterranean Biology 34: 61–98 (2020)
doi: 10.3897/subtbiol.34.49586
http://subtbiol.pensoft.net
Copyright Zegang Feng et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
RESEARCH ARTICLE
Subterranean
Biology Published by
The International Society
for Subterranean Biology
A peer-reviewed open-access journal
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
62
Introduction
Biospeleological studies in the South China Karst (SCK) has rapidly accelerated in re-
cent years. Since 2017, 39 new subterranean-adapted species across several taxonomic
arthropod groups have been described (Gao et al. 2017; Huang et al. 2017; Li and
Wang 2017; Song et al. 2017; Tian et al. 2017, 2018; Deuve and Tian 2018; Li et al.
2019a). Overall, at least 382 cave-dwelling arthropod species are now known from this
region (Ran and Yang 2015; Tian et al. 2016; Li and Wang 2017; Gao et al. 2018;
Feng et al. 2019; Li et al. 2019b; Liu and Wynne 2019). Incidentally, this work has
also resulted in the identication of at least 21 troglomorphic pseudoscorpion species
(refer to Feng et al. 2019; Li et al. 2019b).
In the last 25 years, cave-dwelling pseudoscorpions from China, specically in
Guizhou, Yunnan, Guangxi, Sichuan, and Hubei Provinces, and Beijing and Chongq-
ing Municipalities, total at least 29 pseudoscorpion species (Schawaller 1995; Mahnert
2003, 2009; Mahnert and Li 2016; Gao et al. 2017; Li et al. 2017; Gao et al. 2018;
Feng et al. 2019; Li et al. 2019; Table 1). Of these, 23 are troglobionts and six are
troglophiles (Table 1). With 18 species, Neobisiidae is the most diverse family with
species spanning two genera: Parobisium Chamberlin, 1930 and Bisetocreagris Ćurčić,
1983. Additionally, the families Chernetidae and Chthoniidae contain six and ve
species, respectively.
e pseudoscorpion genus Parobisium was rst established by Chamberlin (1930)
as a subgenus of Neobisium Chamberlin, 1930, and later elevated by Chamberlin and
Malcolm (1960) to generic rank. Parobisium is characterized by the absence of a galea
on the movable cheliceral nger, xed chelal nger with a compact subterminal cluster
of only three tactile setae (et, it, est), and a more diuse subbasal to basal cluster of ve
tactile setae (isb, ist, ib, esb, eb) (Chamberlin 1962). However, for some North Ameri-
can and Asian Parobisium species, the trichobothrium (est) is isolated in the distal half
of the xed nger and has a trichobothrial pattern quite similar to Bisetocreagris.
e key character used to distinguish between these two genera is that Bisetocrea-
gris usually has elongate galeae. Mahnert and Li (2016) intimated the galea is extremely
fragile in Bisetocreagris species. is implies galea may be easily broken or damaged
during collection or transport (Y. Li, pers. com., 18 December 2019). Subsequently,
using galeae as a diagnostic character for describing and identifying species may hinder
accurate classication of this group. In general, Parobisium diers from Bisetocreagris as
there is a distinct and rounded sclerotic knob, rather than an absence of galea (Mori-
kawa 1960; Hong 1996; Mahnert and Li 2016).
During the examination of Guizhou specimens collected by Mingyi Tian (of the
South China Agricultural University, Guangzhou Province) between 2013 and 2017,
we identied several species of Parobisium, which may be undescribed. In most cases,
we had too few specimens to formally describe the species, and in some cases we
had only one specimen. Unfortunately, this can be limiting in describing new spe-
cies, especially given the aforementioned considerations with the galea. To address
this problem, the lead author and colleagues collected additional specimens at the
Cave-dwelling pseudoscorpions of China 63
caves originally sampled by M. Tian. With additional specimens, we were able to both
describe these species and conrm that these Parobisium species have a distinct and
rounded sclerotic knob rather than the absence of galae.
Based upon specimens collected by both M. Tian and the lead author, we describe
four new species of Parobisium from caves in Guizhou Province, China. All species are
subterranean-adapted, and include P. motianense sp. nov., P. qiangzhuang sp. nov., P.
sanlouense sp. nov., and P. tiani sp. nov. We also provide recommendations for manage-
ment of these caves and the cave arthropod communities and habitats they support.
Material and methods
Study area
Guizhou, located in the Yunnan-Guizhou Plateau, is the centrally located province
within the SCK. e karst escarpment within this area is approximately 130,000 km2
encompassing 73% of Guizhou Province (Rong and Yang 2004; He and Li 2016).
Karst formation in Guizhou emerged from a plate group from the Proterozoic to the
Quaternary Period, and consists mainly of shallow marine carbonate deposits (Zhou
et al. 2017). e extensive distribution of carbonate geology and the subtropical mon-
soon climate provide suitable conditions for the development of karst caves. According
to Zhou et al. (2017), Guizhou supports at least 4,960 caves.
e lead author and colleagues searched for and collected pseudoscorpions within
three of the four Guizhou caves (Fig. 1) where M. Tian initially collected specimens.
As none of these caves were subject to previous studies or exploration eorts, cave maps
were not available. We have provided estimations of entrance conguration and cave
length, as well as information on surface vegetation and adjacent human activities.
Motian Cave (Figs 1C, 18) is located ~2 km southwest of Tangbian Town, Ping-
tang County. is limestone cave has one downward sloping oval entrance (~8 meters
high by ~4 meters wide), approximately 2100 meters in length, and extends horizon-
tally. e cave is surrounded by agriculture with the nearest rural residential area less
than 100 m from the cave entrance.
Zharou Cave (Figs 1B, 19) is located ~1 km north of Daying Town, Ziyun County.
is limestone cave has one triangular entrance (~2 meters high and ~3.5 meters wide),
approximately 80 meters in length, and extends horizontally. e surrounding area is
largely disturbed and characterized by low shrubs and weeds (Gramineae); agricultural
elds and a rural residential area are approximately 100 m away.
Sanlou Cave (Figs 1D, 20) is located ~2.5 km northwest of Daoping Town, Fuquan
City. is horizontally-trending cave is approximately 200 meters in length and has an
irregularly round entrance (~4 meters in diameter). Situated near a sand mining opera-
tion, this cave is the primary water source for the village of Daoping. Subsequently, a
reservoir and water delivery system was built in the deepest part of the cave. e cave
has also been designated as a water source protection area.
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
64
Figure 1. Study area, general cave locations, and type locality for each species, Guizhou Province, China.
A Biyun Cave, Parobisium tiani sp. nov. B Zharou Cave, Parobisium qiangzhuang sp. nov. C Motian
Cave, Parobisium motianense sp. nov. D Sanlou Cave, Parobisium sanlouense sp. nov.
Biyun Cave (Figs 1A, 21) is located in Biyun Park, Chengguan Town, Pan County
and is less than 50 m from a rural residential area. is cave has two entrances. One
entrance is dome-shaped (~30 meters wide at the base and 10 m high); during the
rainy season, a river ows into this cave entrance. e second entrance is located about
80 m uphill from the lower entrance and is irregularly round in shape (~30 meters in
diameter). is cave was developed as a tourist cave, and has an unmaintained footpath
paved with concrete, which connects the two entrances.
Field sampling
From 29 July to 5 August 2019, researchers conducted direct intuitive searches (sensu
Wynne et al. 2019) in the estimated deep zone of each cave by examining bat guano,
dead insects, edges of pools and streams, ood detritus, and mud oors (Figs 19B,
20C). Two observers spent approximately two hours searching in Zharou Cave, four
hours searching in Biyun cave, and four observers spent about three hours searching
Sanlou Cave.
Cave-dwelling pseudoscorpions of China 65
Preparation and analysis
Specimens were preserved in 75% ethanol and deposited in the Museum of Hebei
University (MHBU), Baoding, China. Photographs were taken using a Leica M205A
stereomicroscope equipped with a Leica DFC550 camera and LAS software (Ver. 4.6).
We used a Leica M205A stereomicroscope (with a drawing tube) for drawings and
measurements. Chela and chelal hand were measured in ventral view. All measure-
ments are in millimeters (mm) unless noted otherwise. Detailed examination of char-
acters was done using an Olympus BX53 general optical microscope. Temporary slide
mounts were prepared in glycerol.
Terminology
Cave ecosystems typically consist of four environmental zones (Howarth 1980,
1983): (1) entrance zone—or light zone, which represents a combination of sur-
face and cave environmental conditions; (2) twilight zone—occurring slightly deeper
within the cave and has both diminished light conditions and direct inuence of
surface environment; (3) transition zone—aphotic, yet barometric and diurnal shifts
may still occur at a signicantly diminished rate, but the climate is approaching
near stable conditions; and, (4) deep zone—complete darkness, high environmental
stability, near stable temperature, near water-saturated atmosphere, and low to no
airow (usually in the deepest part of the cave). e deep zone represents the region
most conducive to supporting subterranean-adapted animals. Although there are
four primary cave specic functional groups recognized, the species discussed here
have been identied as either troglobionts or troglophiles. Troglobionts are obligate
cave dwellers that require the stable environmental conditions of the deep zone to
complete their life cycle and exhibit morphological characteristics (i.e., troglomor-
phisms) indicative of subterranean adaptation (Sket 2008). Troglophiles (or troglo-
philous organisms) lack troglomorphic characters yet occur facultatively within caves
and complete their life cycles there, but also occur in similar cave-like surface habi-
tats (Barr 1967, Howarth 1983).
Pseudoscorpion terminology and measurements mostly follow Chamberlin (1931)
with some minor modications to the terminology of the trichobothria (Harvey 1992)
and chelicera (Judson 2007). e following abbreviations used for the trichobothria: b
= basal; sb = sub-basal; st = sub-terminal; t = terminal; ib = interior basal; isb = interior
sub-basal; ist = interior sub-terminal; it = interior terminal; eb = exterior basal; esb =
exterior sub-basal; est = exterior sub-terminal; and, et = exterior terminal.
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
66
Table 1. e 29 known cave-dwelling pseudoscorpion species from China. ‘Category’ indicates the func-
tional group in which each species belongs – either troglobiont or troglophile. e number of caves (#
Caves) may be used to infer the level of endemism. Names of administrative provinces where each species
is presently known is also provided.
Taxa Category #Cave Province Reference
Family Chernetidae
Megachernes glandulosus Mahnert, 2009 Troglophile 1Hubei Mahnert (2009)
Megachernes himalayensis (Ellingsen, 1914) Troglophile 1Guangxi Schawaller (1995)
Megachernes tuberosus Mahnert, 2009 Troglophile 1Sichuan Mahnert (2009)
Megachernes vietnamensis Beier, 1967 Troglophile 3Sichuan, Hubei Schawaller (1995)
Nudochernes lipase Mahnert, 2003 Troglophile 1Yunnan Mahnert (2003)
Nudochernes troglobius Mahnert, 2009 Troglophile 2Hubei, Sichuan Mahnert (2009)
Family Chthoniidae
Lagynochthonius bailongtanensis Li, Liu & Shi, 2019 Troglobiont 1Yunnan Li et al. (2019)
Tyrannochthonius akaleus Mahnert, 2009 Troglobiont 1Sichuan Mahnert (2009)
Tyrannochthonius antridraconis Mahnert, 2009 Troglobiont 4Sichuan Mahnert (2009)
Tyrannochthonius chixingi Gao, Wynne & Zhang, 2018 Troglobiont 1Guangxi Gao et al. (2018)
Tyrannochthonius ganshuanensis Mahnert, 2009 Troglobiont 3Sichuan, Hubei Mahnert (2009)
Family Neobisiidae
Bisetocreagris baozinensis Mahnert & Li, 2016 Troglobiont 1Sichuan Mahnert and Li (2016)
Bisetocreagris cavernarum Mahnert & Li, 2016 Troglobiont 1Chongqing Mahnert and Li (2016)
Bisetocreagris chinacavernicola (Schawaller, 1995) Troglobiont 2Sichuan Schawaller (1995)
Bisetocreagris chuanensis Mahnert & Li, 2016 Troglobiont 2Guizhou Mahnert and Li (2016)
Bisetocreagris gracilenta Gao & Zhang, 2017 Troglobiont 1Guizhou Gao et al. (2017)
Bisetocreagris guangshanensis Gao & Zhang, 2017 Troglobiont 1Guizhou Gao et al. (2017)
Bisetocreagris juanxuae Mahnert & Li, 2016 Troglobiont 1Sichuan Mahnert and Li (2016)
Bisetocreagris maomaotou Gao, Wynne & Zhang, 2018 Troglobiont 1Guangxi Gao et al. (2018)
Bisetocreagris martii (Mahnert, 2003) Troglobiont? 1Yunnan Mahnert (2003)
Bisetocreagris scaurum (Mahnert, 2003) Troglobiont 1Yunnan Mahnert (2003)
Bisetocreagris titanium (Mahnert, 2003) Troglobiont 1Yunnan Mahnert (2003)
Bisetocreagris xiaoensis Li & Liu, 2017 Troglobiont 1Yunnan Li et al. (2017)
Parobisium magangensis Feng, Wynne & Zhang, 2019 Troglobiont 1Beijing Feng et al. (2019)
Parobisium motianense sp. nov. Troglobiont 1Guizhou is study
Parobisium qiangzhuang sp. nov. Troglobiont 1Guizhou is study
Parobisium sanlouense sp. nov. Troglobiont 1Guizhou is study
Parobisium tiani sp. nov. Troglobiont 1Guizhou is study
Parobisium yuantongi Feng, Wynne & Zhang, 2019 Troglobiont 1Beijing Feng et al. (2019)
Results
Family Neobisiidae Chamberlin, 1930
Subfamily Neobisiinae Chamberlin 1930
Genus Parobisium Chamberlin 1930
Neobisium (Parobisium) Chamberlin 1930: 17; Beier 1932: 84; Morikawa 1960: 112–
113; Ho 1961: 427.
Parobisium Chamberlin: Chamberlin and Malcolm 1960: 112–113; Chamberlin
1962: 123; Harvey 1991: 394; Mahnert 2003: 744–745.
Type species. Neobisium (Parobisium) magnum Chamberlin, 1930, by original designation.
Cave-dwelling pseudoscorpions of China 67
Key to Parobisium species of China
1 Carapace with eyes or eye spots ......................................................................... 2
– Carapace without eyes or eye spots .................................................................... 5
2 Carapace only with two eyes or eye spots ...................................P. tiani sp. nov.
– Carapace with four eyes or eyespots ..................................................................3
3 Eight setae on posterior margin of carapace; pedipalpal femur 4.65 times longer
than wide, patella 3.14 times longer than wide....................................................
..............................................................P. xiaowutaicum Guo & Zhang, 2016
– Six setae on posterior margin of carapace; both pedipalpal femur and patella more
than 5.7 times longer than wide ........................................................................ 4
4 Pedipalp without granulation; pedipalpal femur 8.91–8.97 times longer than
wide, patella 7.64–7.84 times longer than wide...................................................
...................................................P. magangensis Feng, Wynne & Zhang, 2019
– Pedipalp with granulation present on femur, inside lateral of patella and chelal
hand; pedipalpal femur 6.75 times longer than wide, patella 5.7 times longer than
wide ................................................P. yuantongi Feng, Wynne & Zhang, 2019
5 Carapace with four developed eyes; epistome small, triangular; pedipalp without
granulation .......................................................P. wangae Guo & Zhang, 2016
– Carapace with four eyespots; epistome small, rounded; pedipalp with nely gran-
ulation ..............................................................................................................6
6 Pedipalpal femur 6.50–6.59 times longer than wide, patella 5.07–5.11 times lon-
ger than wide ....................................................................P. sanlouense sp. nov.
– Boyh pedipalpal femur and patella less than 5.0 times longer than wide............7
7 Femur of pedipalp with granulation; pedipalpal femur 3.89–4.11 times longer
than wide, patella 2.54–2.60 times longer than wide ....P. qiangzhuang sp. nov.
– Femur of pedipalp without granulation; pedipalpal femur 4.66–4.9 times longer
than wide, patella 3.09–3.39 times longer than wide ...... P. motianense sp. nov.
Parobisium motianense sp. nov.
http://zoobank.org/D19923FF-CC27-4B00-9A9E-4DB225825490
Figs 2–5
Type material. Holotype male (Ps.-MHBU- GZ17051801): China, Guizhou Prov-
ince, Pingtang County, Tangbian Town, Motian Cave (Figs 1C, 18), [25°38'32.86"N,
104°46'00.36"E], 869 m elevation, 18 May 2017, Mingyi Tian leg. Paratypes: 2 males
(Ps.-MHBU- GZ17051802 & GZ170501803), 1 female (Ps.-MHBU- GZ17051804),
same data as for holotype.
Etymology. Latinized adjective derived from the name of the type locality, Mo-
tian Cave.
Distribution. is species is known only from the type locality.
Diagnosis. Prior to this study, only four species of Parobisium have been reported
in China (Parobisium wangae Guo & Zhang, 2016, Parobisium xiaowutaicum Guo &
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
68
Figure 2. Parobisium motianense sp. nov. Sex indeterminable from photo.
Zhang, 2016, Parobisium magangensis Feng, Wynne & Zhang, 2019 and Parobisium
yuantongi Feng, Wynne & Zhang, 2019). e new troglomorphic species can be dis-
tinguished from other members of the genus Parobisium by following combination
of characters: carapace with four eye spots on a raised surface (P. wangae has four
developed eyes, P. tiani with two faint eye spots; P. magangensis, P. xiaowutaicum and
P. yuantongi lacks eyes/eye spots); epistome small, rounded (small, triangular in P. wan-
gae; triangular, with rounded top in P. tiani and P. yuantongi); carapace with six setae
on posterior margin (eight in P. wangae; eight in P. xiaowutaicum); pedipalpal femur
4.66–4.90 times longer than wide (8.91–8.97 times in P. magangensis; 3.89–4.11 times
in P. qiangzhuang; 6.50–6.59 times in P. sanlouense; 5.63–5.73 times in P. tiani; 3.60–
3.65 times in P. wangae; 6.75 times in P. yuantongi); patella 3.09–3.39 times longer
than wide (7.64–7.84 times in P. magangensis; 2.54–2.60 times in P. qiangzhuang;
5.07–5.11 times in P. sanlouense; 4.52–4.58 times in P. tiani; 1.89–2.16 times in P.
wangae; 5.70 times in P. yuantongi;); pedipalpal hand which is nely granular (smooth
in P. magangensis, P. wangae and P. xiaowutaicum; with granulation present on inside
lateral of femur and chelal hand in P. qiangzhuang and P. sanlouense; with granula-
tion present on femur, inside lateral of patella and chelal hand in P. yuantongi); chela
(with pedicel) 3.72–4.06 times longer than wide (8.67–8.69 times in P. magangensis;
3.12–3.25 times in P. qiangzhuang; 6.08–6.34 times in P. sanlouense; 4.97–5.03 times
in P. tiani; 3.13–3.52 times in P. wangae; 3.14 times in P. xiaowutaicum; 5.70 times in
P. yuantongi); both chelal nger has 95–98 teeth (146–162 in P. magangensis; 69–80 in
Cave-dwelling pseudoscorpions of China 69
Figure 3. Parobisium motianense sp. nov. A Holotype male, dorsal view B Paratype female, dorsal view.
P. qiangzhuang; 119–130 in P. sanlouensis; 57–74 in P. wangae; 73–75 in P. xiaowutai-
cum; 116–118 in P. yuantongi).
Description. Male (Fig. 3A). Carapace, chelicerae, and pedipalps yellowish brown
to reddish brown; abdomen and legs yellowish.
Carapace (Figs 4A, 5A): Smooth, 1.16–1.20 times longer than broad, with a total
of 28–30 setae, including 4 on anterior margin, 6 on posterior margin, and 1 on each
side of anterior lateral margin; with four eye spots on a raised surface (Fig. 4B); epi-
stome small, rounded.
Chelicera (Figs 4C, 5B): Hand with 7 setae, movable nger with 1 submedial
seta; xed nger with 12–14 teeth; movable nger with 14–15 teeth; serrula exterior
with 40–44 lamellae; serrula interior with 25–29 lamellae. Galea (Fig. 5E) replaced by
a small rounded transparent sclerotic knob. Rallum (Fig. 5C) with 8 pinnate blade,
distal-most blade with expanded base, and together with the second blade slightly
separated from the others, proximal one short.
Pedipalps (Figs 4D–E, 5H–J): Apex of coxa rounded, with 5 setae on each side,
pedipalpal coxa with 7 setae. Pedipalp smooth and slender except for hand, which is
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
70
Figure 4. Parobisium motianense sp. nov., holotype male (A–F, H–I), female (G). A Carapace, dorsal
view B Eye area, lateral view C Left chelicera, dorsal view D Right chela, lateral view E Right pedipalp,
dorsal view F Male genitalia G Female genitalia H Left leg I, lateral view I Left leg IV, lateral view.
nely granular. Trochanter 2.04–2.22 times longer than wide, femur 4.66–4.90, pa-
tella 3.09–3.39 times longer than wide, pedicel about half the entire length of patella,
chela (with pedicel) 3.72–4.06, chela (without pedicel) 3.35–3.71 times longer than
wide, movable nger 1.43–1.45 times longer than hand (without pedicel). Fixed che-
lal nger with 8 trichobothria, movable nger with 4, eb and esb on lateral margin of
hand; ib, ist, and isb closely grouped at the base of the xed nger; est slightly distal of
nger middle; it closer to ngertip than et; on movable nger, st nearer to t than to sb,
the distance between sb and b is somewhat equal to that of sb and st (Figs 4D, 5H–I).
Venom apparatus present only in xed chelal nger, venom duct short, not extending
Cave-dwelling pseudoscorpions of China 71
Figure 5. Parobisium motianense sp. nov., holotype male (A–F, H–L), female (G). A Carapace, dorsal
view B Left chelicera, dorsal view C Rallum D Subterminal tarsal seta E Movable nger of chelicera, show-
ing sclerotic knob F Male genitalia G Female genitalia H Right chela, dorsal view I Right chelal ngers,
lateral view J Right pedipalp, dorsal view (trochanter, femur, and patella) K Left leg I, lateral view L Left
leg IV, lateral view. Scale bars: 0.1 mm (C, D–E), 0.25 mm (B, F–G), 0.5 mm (A, I), 1 mm (H, J–L).
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
72
beyond half the distance to et. Fixed chelal nger with 96–98 teeth, movable nger
with 95–97 teeth.
Abdomen: Pleural membrane granulated. Tergal chaetotaxy (I–XI): 8–11/9–
10/9/9–11/10–11/10–11/9–12/11–12/11–12/10–11/6–8; sternal chaetotaxy (IV–
XI): 9–10/14–16/15–16/15/13–16/12–14/12–15/2; stigmata with 5–6 setae; anal
cone with 2 dorsal and 2 ventral setae. Male genital area (Figs 4F, 5F): sternite II with
35–38 scattered setae; sternite III with anteromedian groove anked by one small seta
on each side, with 20–26 posterior setae.
Legs: Coxa chaetotaxy (I–IV): 8–9/6–7/3–4/9–10. Leg I (Figs 4H, 5K): femur
4.37–4.96, patella 2.92–3.22, tibia 6.00–7.27, basitarsus 3.33–3.85, telotarsus 5.50–
5.57 times longer than deep, femur 1.55–1.61 times longer than patella, telotarsus
1.54–1.56 times longer than basitarsus. Leg IV (Figs 4I, 5L): femur + patella 4.36–
5.00 times longer than deep, femur shorter than patella; tibia 8.04–9.05, basitarsus
3.81–3.88, telotarsus 5.44–6.36 times longer than deep, telotarsus 1.35–1.43 times
longer than basitarsus; tibia with one tactile setae (TS=0.52–0.54), basitarsus with
one tactile setae (TS=0.15–0.16), telotarsus with one tactile setae (TS=0.49–0.53);
subterminal tarsal seta (Fig. 5D) bifurcate; arolium not divided, shorter than the slen-
der and simple claws.
Female (paratype; Fig. 3B): Mostly same as holotype.
Chelicera. Hand with 7 setae, movable nger with 1 submedial seta; xed nger
with 15–17 teeth; movable nger with 14–15 teeth; serrula exterior with 41 lamellae;
serrula interior with 23 lamellae. Galea replaced by a conspicuous semicircular trans-
parent sclerotic knob; rallum of 8 blades, similar to holotype.
Pedipalps. Pedipalpal coxa with 8–9 setae. Trochanter 2.08, femur 4.52, patella
3.02, chela (with pedicel) 3.69, chela (without pedicel) 3.33 times longer than wide,
movable nger 1.22 times longer than hand (without pedicel). Fixed chelal nger with
97 teeth, movable nger with 95 teeth.
Abdomen. Tergal chaetotaxy (I–XI): 8/7/9/10/10/11/11/11/11/10/6; sternal chae-
totaxy (IV–XI): 11/16/17/16/16/14/12/2. Female genital area (Figs 4G, 5G): sternite II
with 6–7 setae on each side; sternite III with a row of 20 setae on the posterior margin.
Measurements: (length/breadth or depth in mm; ratios for most characters in pa-
rentheses). Male (holotype and paratypes). Body length 4.26–4.78. Carapace 1.16–
1.20 (1.31–1.36/1.13). Pedipalpal trochanter 2.04–2.22 (0.91–0.96/0.41–0.47),
femur 4.66–4.90 (1.96–2.05/0.40–0.44), patella 3.09–3.39 (1.66–1.73/0.49–0.56),
chela (with pedicel) 3.72–4.06 (3.09–3.13/0.77–0.83), chela (without pedicel)
3.35–3.71 (2.78–2.86/0.77–0.83), hand length (without pedicel) 1.30–1.31, mov-
able nger length 1.86–1.90 (1.43–1.45 times longer than hand without pedicel). Leg
I: trochanter 1.36–1.50 (0.39/0.26–0.28), femur 4.37–4.96 (1.18–1.19/0.24–0.27),
patella 2.92–3.22 (0.74–0.76/0.23–0.26), tibia 6.00–7.27 (1.02–1.09/0.15–0.17),
basitarsus 3.33–3.85 (0.50/0.13–0.15), telotarsus 5.57–5.50 (0.77–0.78/0.14). Leg
IV: trochanter 2.13–2.44 (0.64–0.67/0.27–0.30), femur + patella 4.36–5.00 (1.90–
1.92/0.38–0.44), tibia 8.04–9.05 (1.85–1.90/0.21–0.23), basitarsus 3.81–3.88
(0.61–0.66/0.16–17), telotarsus 5.44–6.36 (0.87–0.89/0.14–0.16).
Cave-dwelling pseudoscorpions of China 73
Female (paratype). Body length 5.99. Carapace 1.14 (1.46/1.28). Pedipalpal tro-
chanter 2.08 (1.02/0.49), femur 4.52 (2.08/0.46), patella 3.02 (1.81/0.60), chela
(with pedicel) 3.69 (3.28/0.89), chela (without pedicel) 3.33 (2.96/0.89), hand length
(without pedicel) 1.47, movable nger length 1.79 (1.22 times longer than hand
without pedicel). Leg I: trochanter 1.47 (0.44/0.30), femur 4.62 (1.20/0.26), patella
3.57 (0.82/0.23), tibia 6.76 (1.15/0.17), basitarsus 3.40 (0.51/0.15), telotarsus 5.13
(0.77/0.15). Leg IV: trochanter 2.28 (0.73/0.32), femur + patella 5.10 (1.99/0.39),
tibia 8.58 (2.06/0.24), basitarsus 3.72 (0.67/0.18), telotarsus 5.29 (0.90/0.17).
Parobisium qiangzhuang sp. nov.
http://zoobank.org/F122663A-2AF0-437C-83C5-791E9739854C
Figs 6–9
Type material. Holotype male (Ps.-MHBU- GZ19080301): China, Guizhou
Province, Anshun City, Ziyun County, Daying Town, Zharou Cave (Figs 1B, 19),
[25°29'24.87"N, 106°18'28.65"E], estimated cave deep zone, 1139 m elevation,
3 August 2019, Zegang Feng, Chen Zhang leg. Paratypes: 2 Males (Ps.-MHBU-
GZ19080302- GZ19080303), same data as for holotype; 1 Female (Ps.-MHBU-
GZ19061201), same location as holotype, 12 June 2018, Sunbin Huang, Zhuanghui
Qin, Mengzhen Chen, Lei Tao leg.
Etymology. e species name, qiangzhuang, was derived from the Latinized Manda-
rin phrase for “strong and hardy” qiáng zhuàng (强壮), which refers to the shape of chela.
Distribution. Species known only from the type locality.
Diagnosis. e subterranean-adapted Parobisium qiangzhuang can be distin-
guished from other members of the genus Parobisium by following combination of
characters: carapace with four eye spots on a raised surface (P. wangae has four devel-
oped eyes, P. tiani with two faint eye spots; P. magangensis, P. xiaowutaicum and P. yu-
antongi lacks eyes/eye spots); epistome small, rounded (small, triangular in P. wangae;
triangular, with rounded top in P. tiani and P. yuantongi); pedipalpal femur 3.89–4.11
times longer than wide (8.91–8.97 times in P. magangensis; 4.66–4.90 times in P. mo-
tianense; 6.50–6.59 times in P. sanlouense; 5.63–5.73 times in P. tiani; 4.65 times in P.
xiaowutaicum; 6.75 times in P. yuantongi); patella 2.54–2.60 times longer than wide
(7.64–7.84 times in P. magangensis; 3.09–3.39 times in P. motianense; 5.07–5.11 times
in P. sanlouense; 4.52–4.58 times in P. tiani; 3.14 times in P. xiaowutaicum; 5.70 times
in P. yuantongi;); pedipalpal hand and inside lateral of femur, which is nely granular
(smooth in P. magangensis, P. wangae and P. xiaowutaicum; with granulation present on
chelal hand in P. tiani; with granulation present on femur, inside lateral of patella and
chelal hand in P. yuantongi); chela (with pedicel) 3.12–3.52 times longer than wide
(8.67–8.69 times in P. magangensis; 3.72–4.06 times in P. motianense; 6.08–6.34 times
in P. qiangzhuang; 4.97–5.03 times in P. tiani; 5.70 times in P. yuantongi); both chelal
nger has 69–80 teeth (146–162 in P. magangensis; 96–98 in P. motianense; 119–130
in P. sanlouensis; 104–112 in P. tiani; 116–118 in P. yuantongi).
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
74
Figure 6. Parobisium qiangzhuang sp. nov. Male habitus.
Description. Male (Fig. 7A). Carapace, chelicerae, and pedipalps reddish brown;
abdomen and legs yellowish.
Carapace (Figs 8A, 9A): Smooth, 1.15–1.24 times longer than broad, with a total
of 28–31 setae, including 4 on anterior margin, 6–8 on posterior margin, and 1–2
on each side of anterior lateral margin; with 4 eye spots on a raised surface (Fig. 8B);
epistome small and rounded.
Chelicera (Figs 8C, 9B): Hand with 7 setae, movable nger with 1 submedial
seta; xed nger with 9–11 teeth; movable nger with 10–13 teeth; serrula exterior
with 34–38 lamellae; serrula interior with 20–24 lamellae. Galea (Fig. 9D) replaced
by a small rounded transparent sclerotic knob. Rallum (Fig. 9C) with 8 pinnate blade,
distal-most blade with expanded base, proximal one short.
Pedipalps (Figs 8D–E, 9H–J): Apex of coxa rounded, with 5 setae on each side,
pedipalpal coxa with 9 setae. Pedipalp smooth except for hand and inside lateral of
femur, which is nely granular. Trochanter 1.84–1.97 times longer than wide, femur
3.89–4.11, patella 2.54–2.60 times longer than wide, pedicel about half the entire
length of patella, chela (with pedicel) 3.12–3.25, chela (without pedicel) 2.86–2.97
times longer than wide, movable nger 1.11–1.15 times longer than hand (without
pedicel). Fixed chelal nger with 8 trichobothria, movable nger with 4, eb and esb on
lateral margin of hand; ib, ist and isb closely grouped at the base of the xed nger;
est slightly distal of nger middle; it-et at same level near ngertip; on movable nger
st nearer to t than to sb, the distance between sb and b is somewhat equal to that of sb
and st (Figs 8D, 9H–I). Venom apparatus present only in xed chelal nger, venom
Cave-dwelling pseudoscorpions of China 75
Figure 7. Parobisium qiangzhuang sp. nov. A Holotype male, dorsal view B Paratype female, dorsal view.
duct short, not extending past half of the distance to et. Fixed chelal nger with 71–75
teeth, movable nger with 69–80 teeth.
Abdomen: Pleural membrane granulated. Tergal chaetotaxy (I–XI): 10–11/11/11–
12/12/12–13/12/12/12/12/12–13/12/7–9; sternal chaetotaxy (IV–XI): 9–11/13–
15/14–16/13–16/13–15/12–14/12–15/3–4; stigmata with 4–5 setae; anal cone with
2 dorsal and 2 ventral setae. Male genital area (Figs 8F, 9E): sternite II with 28–30
scattered setae; sternite III with anteromedian groove anked by one small seta on each
side, with 17–19 posterior setae.
Legs: Coxa chaetotaxy (I–IV): 9–11/7–10/4–5/9–11. Leg I (Figs 8H, 9K): femur
3.52–3.55, patella 2.89–2.94, tibia 5.85–6.67, basitarsus 2.67–2.82, telotarsus 4.42–
4.82 times longer than deep, femur 1.40–4.42 times longer than patella, telotarsus
1.66–1.71 times longer than basitarsus. Leg IV (Figs 8I, 9L): femur + patella 3.63–
3.65 times longer than deep, femur shorter than patella, tibia 6.45–6.79, basitarsus
2.93, telotarsus 4.40–4.57 times longer than deep, telotarsus 1.50–1.56 times longer
than basitarsus; tibia with one tactile setae (TS=0.48–0.50), basitarsus with one tactile
setae (TS=0.11–0.15), telotarsus with one tactile setae (TS=0.41–0.45); subterminal
tarsal seta (Fig. 9G) bifurcate, both branches dentate; arolium not divided, shorter
than the slender and simple claws.
Female (paratype) (Fig. 7B): Mostly same as holotype.
Chelicera. Hand with 7 setae, movable nger with 1 submedial seta; xed nger
with 13 teeth; movable nger with 12 teeth; serrula exterior with 38 lamellae; serrula
interior with 24 lamellae. Galea replaced by conspicuous semicircular transparent scle-
rotic knob; rallum of 8 blades, but similar to holotype.
Pedipalps. Pedipalpal coxa with 10–11 setae. Trochanter 2.02, femur 3.60, patella
2.18, chela (with pedicel) 2.86, chela (without pedicel) 2.63 times longer than wide,
movable nger 1.01 times longer than hand (without pedicel). Fixed chelal nger with
about 72 teeth, movable nger with about 78 teeth.
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
76
Figure 8. Parobisium qiangzhuang sp. nov. holotype male (A–F, H–I), female (G): A Carapace, dorsal
view B Eye area, lateral view C Right chelicera, dorsal view D Right chela, lateral view E Right pedipalp,
dorsal view F Male genitalia G Female genitalia H Right leg I, lateral view I Right leg IV, lateral view.
Cave-dwelling pseudoscorpions of China 77
Figure 9. Parobisium qiangzhuang sp. nov., holotype male (A–E, G–L), female (F). A Carapace, dorsal
view B Right chelicera, dorsal view C Rallum D Movable nger of chelicera, showing sclerotic knob EMale
genitalia F Female genitalia G Subterminal tarsal seta H Right chela, dorsal view I Right chelal ngers,
lateral view J Right pedipalp, dorsal view (trochanter, femur, and patella) K Right leg I, lateral view L Right
leg IV, lateral view. Scale bars: 0.1 mm (C–D, G), 0.25 mm (B, E–F), 0.5 mm (A, K–L), 1 mm (H–J).
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
78
Abdomen. Tergal chaetotaxy (I–XI): 8/11/13/13/12/12/13/13/13/12/5; sternal
chaetotaxy (IV–XI): 7/16/14/14/15/14/12/5. Female genital area (Figs 8G, 9F): sternite
II with 4 setae on each side; sternite III with a row of 12 setae on the posterior margin.
Measurements: (length/breadth or depth in mm; ratios for most characters in pa-
rentheses). Male (holotype and paratypes). Body length 3.61–4.42. Carapace 1.15–
1.24 (1.12–1.18/0.95–0.97). Pedipalpal trochanter 1.84–1.97 (0.70–0.75/0.38),
femur 3.89–4.11 (1.40–1.48/0.36), patella 2.54–2.60 (1.22–1.30/0.48–0.50), chela
(with pedicel) 3.12–3.25 (2.18–2.28/0.67–0.73), chela (without pedicel) 2.86–2.97
(1.99–2.09/0.67–0.73), hand length (without pedicel) 1.02–1.11, movable nger
length 1.17–1.23 (1.11–1.15 times longer than hand without pedicel). Leg I: tro-
chanter 1.17–1.30 (0.27–0.30/0.23), femur 3.52–3.55 (0.74–0.78/0.21–0.22), pa-
tella 2.89–2.94 (0.53–0.55/0.18–0.19), tibia 5.85–6.67 (0.76–0.80/0.12–0.13), ba-
sitarsus 2.67–2.82 (0.31–0.32/0.11–0.12), telotarsus 4.42–4.82 (0.53/0.11–0.12).
Leg IV: trochanter 2.08–2.30 (0.50–0.53/0.23–0.24), femur + patella 3.63–3.65
(1.24–1.27/0.34–0.35), tibia 2.93 (0.41–0.44/0.14–0.15), basitarsus 3.81–3.88
(0.61–0.66/0.16–17), telotarsus 4.40–4.57 (0.64–0.66/0.14–0.15).
Female (paratype). Body length 5.49. Carapace 1.24 (1.50/1.21). Pedipalpal tro-
chanter 2.02 (0.91/0.45), femur 3.60 (1.69/0.47), patella 2.18 (1.46/0.67), chela
(with pedicel) 2.86 (2.69/0.94), chela (without pedicel) 2.63 (2.47/0.94), hand length
(without pedicel) 1.37, movable nger length 1.38 (1.01 times longer than hand
without pedicel). Leg I: trochanter 1.26 (0.34/0.27), femur 3.54 (0.85/0.24), patella
2.86 (0.63/0.22), tibia 7.08 (0.92/0.13), basitarsus 3.17 (0.38/0.12), telotarsus 4.46
(0.58/0.13). Leg IV: trochanter 2.07 (0.60/0.29), femur + patella 4.11 (1.56/0.38),
tibia 7.55 (1.51/0.20), basitarsus 2.81 (0.45/0.16), telotarsus 4.67 (0.70/0.15).
Parobisium sanlouense sp. nov.
http://zoobank.org/EC06FD6A-41EB-47A5-A31A-BC2EC89941A7
Figs 10–13
Type material. Holotype male (Ps.-MHBU-GZ15050201): China, Guizhou Prov-
ince, Fuquan County, Sanlou Cave (Figs 1D, 20), [26°56'46"N, 107°18'47"E],
1280 m elevation, 02 May 2015, Mingyi Tian leg. Paratypes: 3 males (Ps.-MHBU-
GZ19072901, GZ19072902, GZ19072903), 3 females (Ps.-MHBU- GZ19072904,
GZ19072905, GZ19072906), same location as holotype, estimated cave deep zone,
29 July 2019, Zegang Feng, Chen Zhang, Zhaoyi Li, Yonghao Li leg.
Etymology. Latinized adjective derived from the name of the type locality, Sanlou Cave.
Distribution. is species is known only from the type locality.
Diagnosis. is new species can be easily distinguished from other members of the
genus Parobisium by following combination of characters: carapace with four eye spots
on a slightly raised (P. wangae has four developed eyes, P. tiani with two faint eye spots;
P. magangensis, P. xiaowutaicum and P. yuantongi lacks eyes/eye spots); epistome small,
rounded (small, triangular in P. wangae; triangular, with rounded top in P. tiani and P.
yuantongi); pedipalpal femur 6.50–6.59 times longer than wide (8.91–8.97 times in
Cave-dwelling pseudoscorpions of China 79
P. magangensis; 4.66–4.90 times in P. motianense; 3.89–4.11 times in P. qiangzhuang;
5.63–5.73 times in P. tiani; 3.60–3.65 times in P. wangae; 4.65 times in P. xiaowutai-
cum); patella 5.07–5.11 times longer than wide (7.64–7.84 times in P. magangensis;
3.09–3.39 times in P. motianense; 2.54–2.60 times in P. qiangzhuang; 4.52–4.58 times
in P. tiani; 1.89–2.16 times in P. wangae; 3.14 times in P. xiaowutaicum; 5.70 times
in P. yuantongi;); pedipalpal hand and inside lateral of femur, which is nely granular
(smooth in P. magangensis, P. wangae and P. xiaowutaicum; with granulation present on
chelal hand in P. tiani; with granulation present on femur, inside lateral of patella and
chelal hand in P. yuantongi); chela (with pedicel) 6.08–6.34 times longer than wide
(8.67–8.69 times in P. magangensis; 3.72–4.06 times in P. motianense; 3.12–3.25 times
in P. qiangzhuang; 4.97–5.03 times in P. tiani; 3.13–3.52 times in P. wangae; 3.14
times in P. xiaowutaicum; 5.70 times in P. yuantongi); both chelal nger has 119–130
teeth (146–162 in P. magangensis; 95–98 in P. motianense; 69–80 in P. qiangzhuang;
57–74 in P. wangae; 73–75 in P. xiaowutaicum).
Description. Male (Fig. 11A). Carapace, chelicerae, and pedipalps reddish brown
or yellowish brown; abdomen and legs yellowish.
Carapace (Figs 12A, 13A): Smooth, 1.21–1.30 times longer than broad, with a
total of 29–31 setae, including 4 on anterior margin, 7 on posterior margin, and 1–2
on each side of anterior lateral margin; with 4 eye spots on a slightly raised surface (Fig.
12B); epistome small, rounded.
Chelicera (Figs 12C, 13B): Hand with 7 setae, movable nger with 1 submedial
seta; xed nger with 13–14 teeth; movable nger with 13–15 teeth; serrula exterior
with 39–40 lamellae; serrula interior with 25–27 lamellae. Galea (Fig. 13F) replaced
Figure 10. Parobisium sanlouense sp. nov. Male habitus.
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
80
Figure 11. Parobisium sanlouense sp. nov. A Holotype male, dorsal view B Paratype female, dorsal view.
by a small rounded transparent sclerotic knob. Rallum (Fig. 13C) with 8 pinnate blade,
distal-most blade with expanded base, proximal one short.
Pedipalps (Figs 12D–E, 13H, J–K): Apex of coxa rounded, with 5 setae on each
side, pedipalpal coxa with 8–10 setae. Pedipalp smooth and slender except for hand
and inside lateral of femur, which is nely granular. Trochanter 2.86–2.92 times longer
than wide, femur 6.50–6.59, patella 5.07–5.11 times longer than wide, pedicel about
half the entire length of patella, chela (with pedicel) 6.08–6.34, chela (without pedicel)
5.44–5.62 times longer than wide, movable nger 1.51–1.57 times longer than hand
(without pedicel). Fixed chelal nger with 8 trichobothria, movable nger with 4, eb
and esb on lateral margin of hand; ib, ist and isb closely grouped at the base of the xed
nger; est slightly distal of nger at middle; it slightly closer to ngertip than et; on
movable nger st nearer to t than to sb, the latter slightly nearer b than to st (Figs 12D,
13H, J). Venom apparatus present only in xed chelal nger, venom duct short, not
extending past half of the distance to et. Fixed chelal nger with 126–130 teeth, mov-
able nger with 119–126 teeth.
Abdomen: Pleural membrane granulated. Tergal chaetotaxy (I–XI): 7/7–9/9–
10/10/8–10/11/11/11/11–12/11/7; sternal chaetotaxy (IV–XI): 9–10/13/12/12–13/10–
13/13–14/12/3; stigmata with 4–6 setae; anal cone with 2 dorsal and 2 ventral setae.
Male genital area (Figs 12F, 13D): sternite II with 34–36 scattered setae; sternite III with
anteromedian groove anked by one small seta on each side, with 21–30 posterior setae.
Legs: Coxa chaetotaxy (I–IV): 8–9/5–6/5/9. Leg I (Figs 12H, 13L): femur 5.08–
5.73, patella 3.82–4.05, tibia 8.21–8.57, basitarsus 4.00–4.33, telotarsus 6.08–6.58
times longer than deep, femur 1.45–1.48 times longer than patella, telotarsus 1.40–
Cave-dwelling pseudoscorpions of China 81
Figure 12. Parobisium sanlouense sp. nov., holotype male (A–F, H –I), female (G). A Carapace, dorsal
view B Eye area, lateral view C Right chelicera, dorsal view D Right chela, lateral view E Right pedipalp,
dorsal view F Male genitalia G Female genitalia H Right leg I, lateral view I Right leg IV, lateral view.
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
82
1.52 times longer than basitarsus. Leg IV (Figs 12I, 13M): femur + patella 5.22–5.97
times longer than deep, femur shorter than patella, tibia 8.95–9.10, basitarsus 4.06–
4.19, telotarsus 6.93–6.86 times longer than deep, telotarsus 1.43–1.49 times longer
basitarsus; basitarsus with one tactile setae (TS=0.12–0.13), telotarsus with one tactile
setae (TS=0.43–0.52); subterminal tarsal seta (Fig. 13I) bifurcate, both branches den-
tate; arolium not divided, shorter than the slender and simple claws.
Female (paratypes) (Fig. 11B): Mostly same as holotype.
Chelicera. Hand with 7 setae, movable nger with 1 submedial seta; xed nger
with 13–16 teeth; movable nger with 14–15 teeth; serrula exterior with 37–41 lamel-
lae; serrula interior with 23–27 lamellae. Galea (Fig. 13G) replaced by conspicuous sem-
icircular transparent sclerotic knob; rallum of 8–10 blades, similar to that of holotype.
Pedipalps. Pedipalpal coxa with 8–9 setae. Trochanter 2.59–2.89, femur 6.03–6.60,
patella 4.52–4.72, chela (with pedicel) 5.07–5.35, chela (without pedicel) 4.52–4.77
times longer than wide, movable nger 2.00–2.02 times longer than hand (without
pedicel). Fixed chelal nger with 119–128 teeth, movable nger with 115–125 teeth.
Abdomen. Tergal chaetotaxy (I–XI): 6–9/8–11/10–11/9–11/9–10/10–11/11/10–
11/10–11/9–11/7; sternal chaetotaxy (IV–XI): 8–9/12–13/12/12/12–14/12–14/12–
13/3–4. Female genital area (Figs 12G, 13E): sternite II with 4–6 setae on each side;
sternite III with a row of 12–15 setae on the posterior margin.
Measurements: (length/breadth or depth in mm; ratios for most characters in pa-
rentheses). Male (holotype and paratypes). Body length 4.00–4.79. Carapace 1.21–
1.30 (1.32–1.42/1.09–1.09). Pedipalpal trochanter 2.86–2.92 (1.03–1.08/0.36–
0.37), femur 6.50–6.59 (2.34–2.44/0.36–0.37), patella 5.07–5.11 (2.23–2.25/0.44),
chela (with pedicel) 6.08–6.34 (3.59–3.68/0.58–0.59), chela (without pedicel)
5.44–5.62 (3.21–3.26/0.58–0.59), hand length (without pedicel) 1.34–1.38, mov-
able nger length 2.08–2.11 (1.51–1.57 times longer than hand without pedicel). Leg
I: trochanter 1.54–1.57 (0.43–0.44/0.28), femur 5.08–5.73 (1.22–1.26/0.22–1.24),
patella 3.82–4.05 (0.84–0.85/0.21–0.22), tibia 8.21–8.57 (1.15–1.20/0.14), basi-
tarsus 4.00–4.33 (0.52/0.12–0.13), telotarsus 6.08–6.58 (0.73–0.79/0.12). Leg IV:
trochanter 2.41–2.55 (0.70–0.74/0.29), femur + patella 5.22–5.97 (1.93–2.09/0.35–
0.37), tibia 8.95–9.10 (1.88–1.91/0.21), basitarsus 4.06–4.19 (0.65–0.67/0.16), telo-
tarsus 6.93–6.86 (0.96–0.97/0.14).
Female (paratypes). Body length 4.94–6.00. Carapace 1.36–1.39 (1.51–
1.54/1.11). Pedipalpal trochanter 2.59–2.89 (1.01–1.07/0.37–0.39), femur 6.03–
6.60 (2.17–2.31/0.36–0.35), patella 4.52–4.72 (1.99–2.17/0.44–0.46), chela (with
pedicel) 5.07–5.35 (3.40–3.48/0.65–0.67), chela (without pedicel) 4.52–4.77 (3.03–
3.10/0.65–0.67), hand length (without pedicel) 1.31–1.34, movable nger length
1.87–2.00 (1.40–1.53 times longer than hand without pedicel). Leg I: trochanter
1.48–1.56 (0.40–0.42/0.27), femur 5.09–5.48 (1.12–1.26/0.22–0.23), patella 3.90–
4.05 (0.78–0.81/0.20), tibia 7.20–8.14 (1.08–1.14/0.14–0.15), basitarsus 3.69–4.00
(0.48–0.52/0.13), telotarsus 5.21–6.08 (0.73–0.79/0.13–0.14). Leg IV: trochanter
2.10–2.64 (0.61–0.74/0.28–0.29), femur + patella 4.87–5.72 (1.90–2.06/0.36–0.39),
tibia 8.48–9.05 (1.78–1.99/0.21–0.22), basitarsus 3.88–4.19 (0.66–0.67/0.16–0.17),
telotarsus 6.50–7.07 (0.91–0.99/0.14).
Cave-dwelling pseudoscorpions of China 83
Figure 13. Parobisium sanlouense sp. nov., holotype male (A–D, G–M), female (E–F). A Carapace, dor-
sal view B Right chelicera, dorsal view C Rallum D Male genitalia E Female genitalia F Movable nger
of chelicera (male), showing sclerotic knob G Movable nger of chelicera (female), showing sclerotic knob
H Right pedipalp, dorsal view (trochanter, femur, and patella) I Subterminal tarsal seta J Right chelal n-
gers, lateral view K Right chela, dorsal view L Right leg I, lateral view M Right leg IV, lateral view. Scale
bars: 0.5 mm (A–B), 0.1 mm (C, I), 0.25 mm (D–G), 1 mm (H, J–M).
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
84
Parobisium tiani sp. nov.
http://zoobank.org/5E7363D4-BF81-401D-94A2-EA7D654FFBC1
Figs 14–17
Type material. Holotype male (Ps.-MHBU-GZ13070901): China, Guizhou Prov-
ince, Liupanshui City, Pan County, Chengguan Town, Biyun Cave (Figs 1A, 21),
[25°46'29.97"N, 104°38'15.81"E], 1500 m elevation, 9 July 2013, Mingyi Tian leg.
Paratypes: 1 female (Ps.-MHBU-GZ13070902), same location as holotype, 09 July
2013, Mingyi Tian leg; 2 males (Ps.-MHBU-GZ19080501, GZ19080502), 2 females
(Ps.-MHBU-GZ19080503, GZ19080504) same location as holotype, estimated cave
deep zone, 05 August 2019, Zegang Feng, Chen Zhang leg.
Etymology. e name is a patronym to honor Chinese cave biologist, Mingyi
Tian. He provided us with his pseudoscorpion specimens and assisted in developing
this study.
Distribution. is species is known only from the type locality.
Diagnosis. is new troglomorphic species can be easily distinguished from other
members of the genus Parobisium by following combination of characters: carapace with
two faint eye spots (P. wangae has four developed eyes, P. motianense, P. qiangzhuang and
P. sanlouense with four eye spots; P. magangensis, P. xiaowutaicum and P. yuantongi lacks
eyes/eye spots); epistome triangular, with rounded top (small, rounded in P. motianense,
P. qiangzhuang and P. sanlouense; triangular, with rounded top in P. tiani and P. yuanton-
gi; rounded in P. magangensis and P. xiaowutaicum); pedipalpal femur 5.63–5.75 times
longer than wide (8.91–8.97 times in P. magangensis; 4.66–4.90 times in P. motianense;
3.89–4.11 times in P. qiangzhuang; 6.50–6.59 times in P. sanlouense; 3.60–3.65 times in
P. wangae; 4.65 times in P. xiaowutaicum; 6.75 times in P. yuantongi); patella 4.52–4.58
times longer than wide (7.64–7.84 times in P. magangensis; 3.09–3.39 times in P. moti-
anense; 2.54–2.60 times in P. qiangzhuang; 5.07–5.11 times in P. sanlouense; 1.89–2.16
times in P. wangae; 3.14 times in P. xiaowutaicum; 5.70 times in P. yuantongi); pedipal-
pal hand with granulation (smooth in P. magangensis, P. wangae and P. xiaowutaicum;
with granulation present on chelal hand in P. tiani; with granulation present on inside
lateral of femur and chelal hand in P. qiangzhuang and P. sanlouense; with granulation
present on femur, inside lateral of patella and chelal hand in P. yuantongi); chela (with
pedicel) 4.97–5.03 times longer than wide (8.67–8.69 times in P. magangensis; 3.72–
4.06 times in P. motianense; 3.12–3.25 times in P. qiangzhuang; 6.08–6.34 times in P.
sanlouense; 3.13–3.52 times in P. wangae; 3.14 times in P. xiaowutaicum; 5.70 times in
P. yuantongi); both chelal nger has 104–112 teeth (146–162 in P. magangensis; 71–75
in P. qiangzhuang; 57–74 in P. wangae; 73–75 in P. xiaowutaicum).
Description. Male (Fig. 15A). Carapace, chelicerae, and pedipalps reddish brown
or yellowish brown; abdomen and legs yellowish or yellowish brown.
Carapace (Figs 16A, 17A): Smooth, 1.22–1.27 times longer than broad, with a
total of 24 setae, including 4 on anterior margin, 6 on posterior margin, and 1 on
each side of anterior lateral margin; with 2 faint eye spots on a at surface; epistome
triangular, with rounded top.
Cave-dwelling pseudoscorpions of China 85
Figure 14. Parobisium tiani sp. nov. Male habitus.
Chelicera (Figs 16C, 17B): Hand with 7 setae, movable nger with 1 submedial
seta; xed nger with 13–16 teeth; movable nger with 13–16 teeth; serrula exterior
with 39–42 lamellae; serrula interior with 20–24 lamellae. Galea (Fig. 17D) replaced
by a small rounded transparent sclerotic knob. Rallum (Fig. 17C) with 8 pinnate blade,
distal-most blade with expanded base, and together with the second blade separated
from the others, proximal one short.
Pedipalps (Figs 16E–F, 17H–I, K): Apex of coxa rounded, with 5 setae on each
side, pedipalpal coxa with 7–8 setae. Pedipalp smooth and slender except for hand,
which is nely granular. Trochanter 2.67–2.70 times longer than wide, femur 5.63–
5.73, patella 4.52–4.58 times longer than wide, pedicel about half the entire length of
patella, chela (with pedicel) 4.97–5.03, chela (without pedicel) 4.39–4.40 times longer
than wide, movable nger 1.44–1.45 times longer than hand (without pedicel). Fixed
chelal nger with 8 trichobothria, movable nger with 4, eb and esb on lateral margin
of hand; ib, ist and isb closely grouped at the base of the xed nger; est slightly distal
of nger middle; it closer to ngertip than et; on movable nger, st nearer to t than to
sb, the latter slightly nearer st than to b (Figs 16F, 17H–I). Venom apparatus present
only in xed chelal nger, venom duct short, not extending past half of the distance to
et. Fixed chelal nger with 104–109 teeth, movable nger with 105–112 teeth.
Abdomen: Pleural membrane granulated. Tergal chaetotaxy (I–XI): 8–9/ 8–9/ 9/
10/ 9/ 9–11/ 9–10/ 10–11/ 10–12/ 10–11/ 6–7; sternal chaetotaxy (IV–XI): 7–8/
12–13/ 14/ 12–13/ 11–13/ 11–12/ 11–12/ 3–4; stigmata with 5–6 setae; anal cone
with 2 dorsal and 2 ventral setae. Male genital area (Figs 16G, 17F): sternite II with
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
86
Figure 15. Parobisium tiani sp. nov. A Holotype male, dorsal view B Paratype female, dorsal view.
34–43 scattered setae; sternite III with anteromedian groove anked by one small seta
on each side, with 15 posterior setae.
Legs: Coxa chaetotaxy (I–IV): 6–7/ 4–5/ 4–5/ 7. Leg I (Figs 16I, 17L): femur
6.27–6.50, patella 3.54–4.23, tibia 8.87–9.08, basitarsus 3.73–3.93, telotarsus 5.07–
5.85 times longer than deep, femur 1.54–1.62 times longer than patella, telotarsus
1.36–1.38 times longer than basitarsus. Leg IV (Figs 16J, 17M): femur + patella 5.30–
5.46 times longer than deep, femur shorter than patella, tibia 9.04–9.75, basitarsus
4.06–4.41, telotarsus 5.61–6.20 times longer than deep, telotarsus 1.35 times longer
than basitarsus; basitarsus with a tactile setae in basally (TS=0.12–0.15), telotarsus with
a tactile setae in middle (TS=0.46–0.47); subterminal tarsal seta (Fig. 17J) bifurcate,
both branches dentate; arolium not divided, shorter than the slender and simple claws.
Female (paratypes) (Fig. 15B): Mostly same as holotype.
Chelicera. Hand with 7 setae, movable nger with 1 submedial seta; xed nger
with 13–16 teeth; movable nger with 12–19 teeth; serrula exterior with 39–41 lamel-
lae; serrula interior with 22–23 lamellae. Galea (Fig. 17E) replaced by a semicircular
transparent sclerotic knob; rallum of 8–9 blades, similar to holotype.
Pedipalps. Trochanter 2.44–2.61, femur 5.59–5.61, patella 4.38–4.57 times long-
er than wide; chela (with pedicel) 4.25–4.46 times longer than wide, chela (without
pedicel) 3.79–4.00 times longer than wide, movable nger 1.26–1.28 times longer
than hand (without pedicel). Fixed chelal nger with 96–106 teeth, movable nger
with 97–105 teeth.
Abdomen. Tergal chaetotaxy (I–XI): 6–7/ 7–9/ 8–9/ 8–10/ 9–11/ 10–12/ 9–12/
9–12/ 11–12/ 11/ 6–7; sternal chaetotaxy (IV–XI): 8–9/ 13–15/ 13–16/ 13–15/ 13–
Cave-dwelling pseudoscorpions of China 87
Figure 16. Parobisium tiani sp. nov., holotype male (A–C, E–G, I–J), female (D, H). A Carapace,
dorsal view B Eye area, lateral view C Right chelicera of male, dorsal view D Right chelicera of female,
dorsal view E Right pedipalp, dorsal view F Right chela, lateral view G Male genitalia H Female genitalia
I Right leg I, lateral view J Right leg IV, lateral view.
15/ 12–13/ 12–14/ 3. Female genital area (Figs 16H, 17G): sternite II with 3–8 setae
on each side; sternite III with a row of 13–16 setae on the posterior margin.
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
88
Figure 17. Parobisium tiani sp. nov., holotype male (A–D, F, H–M), female (E, G). A Carapace, dorsal
view B Right chelicera, dorsal view C Rallum D Movable nger of chelicera (male), showing sclerotic
knob E Movable nger of chelicera (female), showing sclerotic knob F Male genitalia G Female genitalia
H Right chela, dorsal view I Right chelal ngers, lateral view J Subterminal tarsal seta K Right pedipalp,
dorsal view (trochanter, femur, and patella) L Right leg I, lateral view M Right leg IV, lateral view. Scale
bars: 0.1 mm (C–E, J), 0.25 mm (B, F–G), 0.5 mm (A), 1 mm (H–I, K–M).
Cave-dwelling pseudoscorpions of China 89
Measurements: (length/breadth or depth in mm; ratios for most characters in pa-
rentheses). Male (holotype and paratypes). Body length 3.87–5.09. Carapace 1.22–
1.27 (1.32–1.43/1.08–1.13). Pedipalpal trochanter 2.67–2.70 (1.04–1.16/0.39–
0.43), femur 5.63–5.73 (2.29–2.42/0.40–0.43), patella 4.52–4.58 (2.17–2.38/0.48–
0.52), chela (with pedicel) 4.97–5.03 (3.28–3.52/0.66–0.70), chela (without pedicel)
4.39–4.40 (2.90–3.08/0.66–0.70), hand length (without pedicel) 1.32–1.36, movable
nger length 1.91–1.96 (1.44–1.45 times longer than hand without pedicel). Leg I:
trochanter 1.35–1.50 (0.42/0.28–0.31), femur 6.27–6.50 (1.38–1.43/0.22), patella
3.54–4.23 (0.85–0.93/0.22–0.24), tibia 8.87–9.08 (1.18–1.33/0.13–0.15), basitarsus
3.73–3.93 (0.55–0.56/0.14–0.15), telotarsus 5.07–5.85 (0.76/0.13–0.15). Leg IV:
trochanter 2.43–2.79 (0.68–0.78/0.28), femur + patella 5.30–5.46 (2.13–2.28/0.39–
0.43), tibia 9.04–9.75 (2.17–2.34/0.24), basitarsus 4.06–4.41 (0.69–0.75/0.17), telo-
tarsus 5.61–6.20 (0.93–1.01/0.15–0.18).
Female (paratypes). Body length 4.63–5.48. Carapace 1.19–1.27 (1.30–1.60/1.09–
1.26). Pedipalpal trochanter 2.44–2.61 (0.95–1.15/0.39–0.44), femur 5.59–5.61
(2.18–2.47/0.39–0.44), patella 4.38–4.57 (2.06–2.47/0.47–0.54), chela (with pedi-
cel) 4.25–4.46 (3.03–3.61/0.68–0.85), chela (without pedicel) 3.79–4.00 (2.72–
3.22/0.68–0.85), hand length (without pedicel) 1.29–1.55, movable nger length
1.65–1.95 (1.26–1.28 times longer than hand without pedicel). Leg I: trochanter
1.47–1.60 (0.40–0.44/0.25–0.30), femur 5.39–5.92 (1.24–1.48/0.23–0.25), patella
3.75–4.18 (0.75–0.92/0.20–0.22), tibia 7.00–7.33 (1.05–1.32/0.15–0.18), basitar-
sus 3.14–4.07 (0.44–0.61/0.14–0.15), telotarsus 5.00–5.43 (0.65–0.76/0.13–0.14).
Leg IV: trochanter 2.48–2.61 (0.67–0.81/0.27–0.31), femur + patella 5.91–5.97
(2.01–2.27/0.34–0.38), tibia 9.38–9.67 (1.97–2.32/0.21–0.24), basitarsus 3.82–4.53
(0.65–0.77/0.17), telotarsus 6.00–6.13 (0.90–0.98/0.15–0.16).
Discussion
Our work has increased the number of Chinese cave-dwelling pseudoscorpions from 25
to 29 species. In addition, we found that female individuals may have a distinctly rounded
sclerotic knob, while the sclerotic knob in males was not often obvious. Uncertainty of the
characteristics of the sclerotic knob makes versus the damaged of galeae it dicult to iden-
tify and describe species – especially when only a single-sex specimen is available. We rec-
ommend that future researchers: (1) collect multiple specimens to help ensure both adult
males and females are collected (enabling further examination and study of the sclerotic
knob across additional specimens and species); (2) carefully collect pseudoscorpions to
avoid damaging the galeae; (3) cautiously examine the galeae particularly when specimens
are few; and, (4) use scanning electron microscopy (e.g., Cokendolpher and Krejca 2010)
to more accurately examine the structure of the galeae. e latter will enable researchers to
determine whether galea are reduced or altered during collection or transportation.
As with most Chinese hypogean pseudoscorpions (Table 1; Schawaller 1995;
Mahnert 2003, 2009; Mahnert and Li 2016; Gao et al. 2017; Li et al. 2017; Gao et al.
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
90
Figure 18. Motian Cave, type locality of Parobisium motianense sp. nov. A Surrounding vegetation and
agricultural areas with cave entrance (Red arrow) B Entrance C Inside the cave entrance D Stalactites E
Cave landscape.
2018; Feng et al. 2019; Li et al. 2019), these four new species are currently considered
single cave endemics. However, this may be due to limited investigations in the region,
rather than actual short-range distributions of these species. ere is a high density of
caves in Guizhou – in particular, numerous caves occur within a 5 km radius of the
caves containing these species. Specically, there are at least three caves with a 5 km
radius of Zhaorou Cave, no fewer than seven caves with a 5 km radius of the Sanlou
Cave, and at least 10 caves with a 5 km radius in Biyun Cave. None of these caves have
been inventoried for cave-dwelling arthropods. As a result, it is possible our newly
described species are actually restricted to a geological formation rather than a single
cave (Schawaller 1995; Mahnert 2009; Mahnert and Li 2016; Table 1). Additional
investigations will be required to make this distinction.
While it is becoming increasingly well-established that caves in China are rich
in cave biological resources and support subterranean-adapted species with highly re-
stricted distributional ranges, the county presently lacks policies or a governmental
agency to protect and manage subterranean natural resources. is presents challenges
for conservation and management of sensitive subterranean animal populations. Un-
fortunately, human activities (i.e., urbanization, mining, and other related activities) in
karst areas, and the development of tourist caves, continue without prior evaluations
of the potentially sensitive natural resources and/or the biodiversity they may support
Cave-dwelling pseudoscorpions of China 91
Figure 19. Zharou Cave, type locality of Parobisium qiangzhuang sp. nov. A entrance B Area where P.
qiangzhuang specimens were collected.
(Whitten 2009). erefore, we recommend the distributions of the new species de-
scribed here be more thoroughly established through sampling of the abovementioned
caves surrounding the type localities. Importantly, once we determine whether these
species are single cave or regional endemics, this information may be used to guide
management policies to protect these animals and their habitats.
Local human activities (Figs 20A, D, 21D) are more likely to have signicant
impacts on these cave-dwelling species, which could result in their imperilment or
potentially their extinction. In fact, the four new Guizhou pseudoscorpions and their
habitats are directly threatened by human activities. All four caves were close to hu-
man settlements and/or agricultural areas (within 0 m to 100 m) and were aected to
varying degrees by other human activities. Motian Cave is surrounded by agricultural
activities. For Zharou Cave, agricultural activities are less than 100 m away. In both
cases, pesticide and fertilizer residues may contaminate the caves via runo (Castaño-
Sánchez et al. 2019). Although the entrance of Zharou Cave is somewhat obscured
by vegetation, local residents are aware of this cave, and we observed recent evidence
of human activities. e entrance of the Sanlou Cave is about 50 m from sand min-
ing operations, and the deepest part of the cave has been modied and converted to a
reservoir. Additionally, the sand mining facility and the water extraction activities may
ultimately aect the survival of P. sanlouense sp. nov.
Biyun Cave, located in Biyun county park, is a tourist cave. Evidence of human
activity was observed throughout the cave, which included refuse, remnants of bon-
res, and grati on the cave walls. Subsequently, cave habitats have been damaged to
varying degrees. During our work, we observed at least 10 tourists visiting the cave.
Fortunately, because of the muddy and steep path at the back of the cave, we suspect
fewer visitors will be willing to access the area where we found P. tiani sp. nov. As a
result, this habitat may be somewhat protected.
Extinction is often characterized by time lags, and at-risk populations may persist
for long periods of time near extinction thresholds prior to becoming extinct (e.g.,
Brooks et al. 1999, Hanski and Ovaskainen 2002, Vellend et al. 2006). ese “ex-
Zegang Feng et al. / Subterranean Biology 34: 61–98 (2020)
92
Figure 20. Sanlou Cave, type locality of Parobisium sanlouense sp. nov. A Surrounding surface vegetation
with cave entrance (white arrow) and sand mining operations adjacent to the cave B Entrance C Area
where P. sanlouense specimens were collected D Terminus of Sanlou Cave with the potable water reservoir;
specimens were collected near the reservoir.
tinction debts” (see Tilman et al. 1994) may occur when populations become isolat-
ed following signicant environmental perturbations or intensive human activities.
Both P. sanlouense sp. nov. and P. tiani sp. nov. occur in areas of intensive human ac-
tivities. is further emphasizes the need for additional surveys to determine whether
these species are single cave or regional endemics. is distinction will be of critical
importance in determining the sensitivity of these animals to current human activi-
ties within and at proximity to the caves where they occur. Moreover, understanding
their distributions will be required to develop eective monitoring protocols – if
additional evidence supports that either (or both) species are single cave endemics.
ere are other measures that should be examined to protect sensitive cave-dwell-
ing species and their habitats. An outreach campaign to help educate villagers, school
children, and tourists concerning the vulnerability of cave biological resources should
be considered (refer to Mammola et al. 2019). At Biyun Cave, it may be worth post-
ing educational signs (within the village and perhaps near the entrance) to explain the
sensitivity of cave natural resources and that endemic species occur within, as well as
guidelines for reducing human impacts to sensitive cave resources.
Cave-dwelling pseudoscorpions of China 93
Figure 21. Biyun Cave, type locality of Parobisium tiani sp. nov. A Surrounding surface habitat with two
entrances, entrance 1 (white arrow) and entrance 2 (red arrow) B Entrance 1 (white arrow) C Entrance 2
(red arrow) D Passage where specimens were collected (blue arrow).
As research on cave biological resources in southern China continues, numerous
additional new species with restricted ranges will be described. Understanding their
distributions and the functional roles they play in these often highly sensitive ecologi-
cal communities will be of paramount importance for developing management plans
to protect both sensitive species and their habitat. rough these and other eorts, we
hope our ndings and data collected in the future will be employed to help shape ef-
fective cave resource management in China.
Acknowledgments
We are grateful to Mingyi Tian for contributing specimens and providing us with in-
formation and images (Figs 2, 18), on the caves he sampled. is work was supported
by the National Natural Science Foundation of China (No. 31872198), and the Min-
istry of Science and Technology of the People’s Republic of China (MOST Grant No.
2015FY210300).
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94
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